VIOLACEAE

堇菜科 jin cai ke Chen Yousheng (陈又生)1, Yang Qiner (杨亲二)1; Hideaki Ohba2, Vladimir V. Nikitin3 Herbs annual or perennial, shrubs, or subshrubs, sometimes scandent, rarely small trees. Leaves simple, usually alternate, some- times opposite, with small or leaflike stipules, petiolate, margin entire, serrate, or dissected. Flowers bisexual or unisexual, rarely polygamous, actinomorphic or zygomorphic, solitary or in axillary or terminal, spicate, paniculate, or racemose inflorescences, 2- bracteolate, sometimes cleistogamous. Sepals 5, equal or unequal, imbricate, persistent. Petals 5, imbricate or convolute, unequal, an- terior one usually larger than others, saccate, gibbous or spurred at base. Stamens 5; anthers erect, free or connivent or connate, con- nectives often dilated into membranous appendages; filaments very short or absent, anterior 2 stamens with spurlike nectary at base. Ovary superior, 1-loculed, 3–5-carpelled, syncarpous, with 3–5 parietal placentae each with 1 to many anatropous ovules; style sim- ple; stigmas variously shaped. Fruit a loculicidal capsule, usually with elastic and abaxially carinate valves, rarely baccate. Seeds often carunculate; testa hard, nitid, often with oily bodies, sometimes alate; endosperm copious, fleshy; embryo erect. Twenty-two genera and 900–1000 species: worldwide, but particularly in tropical regions; three genera and 101 species (36 endemic, three introduced) in China. Wang Chingrui. 1991. Violaceae. Fl. Reipubl. Popularis Sin. 51: i–v, 1–148. 1a. Shrubs or trees; inflorescences mostly more than 5-flowered; flowers actinomorphic, not spurred, not cleistogamous ...... 1. Rinorea 1b. Herbs or subshrubs; inflorescences 1–3-flowered; flowers distinctly zygomorphic (lowest petal spurred), or cleistogamous. 2a. Subshrubs; sepals basally not auriculate ...... 2. Hybanthus 2b. Herbs; sepals basally auriculate ...... 3. Viola 1. RINOREA Aublet, Hist. Pl. Guiane, 235. 1775, nom. cons.

三角车属 san jiao che shu Conohoria Aublet. Shrubs or small trees. Leaves alternate, rarely opposite, margin entire or serrate; stipules deciduous. Inflorescences axillary or terminal, racemose, paniculate, or sometimes cymose, rarely flowers solitary. Flowers bisexual, actinomorphic, small. Sepals sub- equal, leathery. Petals equal or subequal, not spurred. Filaments free or ± connate, inserted inside at top of disk; connectives dilated abaxially from base or from apex into thin, scarious appendages exceeding anthers. Disk circular, slightly 5-lobed. Ovary ovoid, with few or numerous ovules; styles erect; stigmas terminal, not divided. Capsule usually 3-valved, rarely 2-valved. Seeds few, ellipsoid, glabrous or pubescent. About 340 species: tropical Africa, tropical America, and tropical Asia; four species (one endemic) in China. 1a. Disk 5-lobed; stigmas slightly 3-lobed; stamens without filaments, connectives with granular scales ...... 4. R. virgata 1b. Disk dilated and circular; stigmas not divided; stamens with filaments, connectives without abaxial scales. 2a. Appendages at top of connectives narrowly triangular, less than 1 mm, anthers lanate at base adaxially, apex barbate; stipules later deciduous; petioles 2–5 mm ...... 3. R. erianthera 2b. Appendages at top of connectives not narrowly triangular, anthers without cushionlike tomentum at base adaxially, apex not barbate; stipules caducous; petioles 7–12 mm. 3a. Leaf veins raised on both surfaces, very conspicuous; pedicels to 1 cm; fruit small, less than 1 cm; seeds with brown dots ...... 1. R. bengalensis 3b. Leaf veins slightly raised abaxially, not very conspicuous; flowers subsessile; fruit large, ca. 2 cm; seeds creamy-white ...... 2. R. longiracemosa 1. Rinorea bengalensis (Wallich) Kuntze, Revis. Gen. Pl. 1: glandulosa Merrill (1928), not Merrill (1917); R. wallichiana 42. 1891. (J. D. Hooker & Thomson) O. Kuntze.

三角车 san jiao che Shrubs or small trees, 1–5 m tall. Young branches with conspicuous leaf scars, greenish, glabrous or with very few Alsodeia bengalensis Wallich, Trans. Med. Soc. Calcutta hairs; old branches dark brown. Leaves alternate, old ones 7: 224. 1835; A. wallichiana J. D. Hooker & Thomson; Rinorea subleathery, slightly nitid; stipules lanceolate, 1–1.7 cm, apex

1 Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 2 University of Tokyo, 7–3–1 Hongo, Bunkyo-ku, Tokyo, Tokyo 113–0033, Japan. 3 V. L. Komarov Botanical Institute, Prof. Popov Street 2, 197376 Saint Petersburg, Russia.

72 VIOLACEAE 73

acuminate, fugacious, scar circular; petiole 5–12 mm, glabrous; clustered, petioles and stipules puberulous when young; stipules leaf blade elliptic-lanceolate or elliptic, (2.5–)5–12(–17) × 1.5– deciduous, narrowly triangular-lanceolate, subequaling petioles; 6 cm, veins raised on both sides, lateral veins alternate, 6–9- petiole very short, 2–5 mm; leaf blade yellow-green, rhombic- paired, veinlets reticulate, dense and conspicuous, base cuneate, elliptic to obovate-elliptic, 1.5–6.5 × 0.6–2.5 cm, membranous, rarely subrounded, margin serrulate, remotely toothed near base base cuneate, margin serrulate, apex acute to shortly acuminate. or subentire, apex acuminate. Flowers white, in axillary con- Flowers 1–3 in cymes, axillary in long branches; pedicels 1–1.5 tracted racemes; peduncles absent; pedicel to 1 cm, slightly cm, slender, hairy, with 2 bracteoles less than 1 mm at middle yellow tomentose. Sepals broadly lanceolate, ca. 2 mm, abaxi- or below. Sepals 5, yellow-green, 4.5–6 × 2–5.5 mm, imbricate; ally yellow-brown tomentose. Petals ovate-oblong, ca. 5 mm, outer 3 suborbicular, densely veined, apex apiculate; inner 2 apex recurved. Stamens with short filaments; anthers 2-loculed, elliptic, inconspicuously veined, hairy abaxially, margin ciliate. oblong, longitudinally dehiscent, appendages at top of con- Petals 5, white, brownish when dry, narrowly lanceolate, 9–11 nectives broadly ovate. Capsule subglobose, 3-valved. Seeds × 1.5–2 mm, hairy above middle abaxially, margin ciliate, re- broadly ovoid, glaucous, with brown dots. Fl. Apr–May and flexed after anthesis. Stamens yellowish white, ca. 9 mm; fila- Oct, fr. Sep. 2n = 16. ments linear, 2.5–3.5 mm; anthers linear, equaling or slightly exceeding filaments, lanate at base adaxially, thecae acuminate Thickets, dense forests; below 600 m. SW Guangxi, Hainan [In- dia, Malaysia, Myanmar, Sri Lanka, Thailand, Vietnam; NW Australia]. at apex, barbate, appendages at top of connectives narrowly tri- angular, less than 1 mm. Ovary narrowly globose, 3–5-angled; 2. Rinorea longiracemosa (Kurz) Craib, Fl. Siam. 1: 90. 1925. styles straight or curved, slender. Seeds (immature) brown, or- 短柄三角车 duan bing san jiao che bicular-reniform, compressed-flat, smooth, not nitid. Fl. Jun– Jul, fr. unknown. Alsodeia longiracemosa Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 39(2): 63. 1870. ● Thickets; ca. 1300 m. SW Sichuan (Jinyang). Shrubs or small trees, 1–2 m tall. Young branches incon- 4. Rinorea virgata (Thwaites) Kuntze, Revis. Gen. Pl. 1: 42. spicuously cicatricate, yellow-brown tomentose; old branches 1891. gray-white, glabrous. Leaves alternate; stipules dark brown, nar- 鳞隔堇 lin ge jin rowly lanceolate or subulate, ca. 4 mm, slightly hairy, fuga- cious; petiole ca. 7 mm, yellow-brown puberulous when young, Scyphellandra virgata Thwaites, Enum. Pl. Zeyl. 21. sparsely puberulous when old; leaf blade elliptic or elliptic- 1858; Alsodeia virgata (Thwaites) J. D. Hooker & Thomson; lanceolate, (7–)12–14(–16) × (1.5–)3.5–4.5(–6) cm, glabrous, Rinorea pierrei (H. Boissieu) Melchior; S. pierrei H. Boissieu. veins not very conspicuous, slightly raised abaxially, midvein Shrubs erect, ca. 1 m tall. Young branches puberulous; old yellow-brown hairy near base, margin remotely serrate, teeth ones gray-white, glabrous. Lower leaves often 2- or 3-clustered, denser in upper half, apex caudate or long acuminate. Flowers upper ones alternate; stipules small, shorter than petioles, lan- glaucous, subsessile, in axillary contracted-racemes. Sepals 5, ceolate or subtriangular, slightly raised abaxially, pubescent; persistent, subtriangular. Corolla campanulate; petals 5, persis- petiole short, ca. 3 mm, tomentose; leaf blade ovate or elliptic, tent, subequal, elliptic. Stamens with short filaments. Capsule highly variable in size, (1.5–)3–4(–5) × (0.8–)1.5–2(–2.5) cm, subglobose, ca. 2 cm. Seeds creamy-white, subglobose. Fl. un- veins slightly raised, slightly puberulous, margin serrulate, apex known, fr. May. obtuse or ± acute. Flowers small, unisexual, actinomorphic, Lowland primary or secondary forests; below 1000 m. Hainan axillary, solitary or clustered. Staminate flowers: pedicels ca. 5 (Lingshui) [Cambodia, Indonesia, Laos, Malaysia, Myanmar, Thailand, mm, slender, slightly hairy or glabrous; sepals 5, subtriangular, Vietnam]. shortly hairy; petals 5, oblong, outside slightly hairy, apex acu- In FRPS (51: 4. 1991), the name Rinorea sessilis (Loureiro) minate; stamens subequaling petals, anthers 2-thecate, longitu- Kuntze was misapplied to this species. dinally dehiscent, connectives yellow, subulate; ovary reduced. Rinorea longiracemosa is very rare in China. Pistillate flowers: pedicels ca. 3 mm, glabrous, with 3 or 4 sub- triangular bracteoles at base; sepals persistent, subtriangular, 3. Rinorea erianthera C. Y. Wu & Chu Ho, Acta Bot. Yun- margin ciliolate; petals oblong, outside slightly hairy; stamens nan. 1(1): 149. 1979. reduced, connectives lanceolate, conspicuous, anthers absent; 毛蕊三角车 mao rui san jiao che ovary ovoid-orbicular, glabrous, styles erect, stigmas capitate, 3-lobed. Capsule oblong, 3-valved, rarely 2-valved, apex cau- Shrubs deciduous, 2–3 m tall. Young branchlets yellow- date. Fl. spring–summer, fr. summer–autumn. brown, puberulous; old ones purplish brown, glabrous, dotlike lenticellate. Leaves on long branches of current year larger and Forest margins, thickets; below 600 m. Hainan [Laos, Myanmar, sparse; leaves on short branches of previous year smaller and Sri Lanka, Thailand, Vietnam]. 2. HYBANTHUS Jacquin, Enum. Syst. Pl. 2, 17. 1760, nom. cons.

鼠鞭草属 shu bian cao shu Small shrubs [or herbs]. Leaves alternate, ± sessile; stipules in pairs, small; leaf blade narrowly elliptic to almost needle-shaped, margin entire to incised. Flowers axillary, bisexual, zygomorphic, solitary, with a pair of bracteoles at joint of peduncle and pedicel.

74 VIOLACEAE

Sepals persistent, ± triangular, nearly equal, margin entire, without appendages at base. Petals unequal, posterior ones straight and small, anterior one extended into a lip, with a short spur at base. Stamens connivent around gynoecium; filaments free, short; anthers 2-celled, with a distinct membranous connective appendage, lower two with a patent gland at base. Ovary: ovules numerous, on 3 parietal placentas; style simple, geniculate at base, conspicuously clavate, with an anterior opening stigmatic cavity. Fruit a 3-valved capsule, subglobose, valves leathery. Seeds numerous, ellipsoid, with longitudinal ribs. About 150 species: chiefly in tropical and subtropical regions of the world; one species in China.

1. Hybanthus enneaspermus (Linnaeus) F. Mueller, Fragm. blue-purple; pedicels not exceeding leaves, 2-bracteolate near 10: 81. 1876. flower; bracteoles opposite, lanceolate or subtriangular, mem- branous, ciliate. Sepals subequal, lanceolate, 2–3 mm, margin 鼠鞭草 shu bian cao membranous, white, ciliate. Petals irregular, upper 2 smallest, oblong, apex acute, lateral 2 ovate-orbicular, apex slightly re- Viola enneasperma Linnaeus, Sp. Pl. 2: 937. 1753; Hy- curved, lower 1 largest, 6–7 mm, ca. 2 × as large as others, an- banthus suffruticosus (Linnaeus) Baillon; Ionidium ennea- terior half broadest, 2 lateral sides incurved, more deeply spermum (Linnaeus) Ventenat; I. suffruticosum (Linnaeus) colored, shortly spurred. Stamens free; filaments short, 2 with Gingins; V. suffruticosa Linnaeus. curved calcarlike (spurlike) appendages; anthers longitudinally Subshrubs, with a long, slender tap root. Stems numerous, dehiscent, appendages at top of connectives yellow-brown, procumbent, usually 10–20 cm tall, rarely to 40 cm. Young membranous. Ovary subglobose; styles clavate, base dilated, branches canaliculate, ± puberulous. Leaves alternate, sessile or slightly curved in upper narrow part; stigmas slightly in- subsessile; stipules greenish, triangular or subulate, very small, curved. Capsule pendulous, globose. Seeds creamy-yellow, ciliate; leaf blade linear-lanceolate, linear-oblanceolate, or nar- ovoid-orbicular, conspicuously longitudinally striate. Fl. Jun– rowly spatulate, unequal in size, 0.5–3.5 × 0.2–0.5 cm, midvein Jul, fr. Jul–Aug. 2n = 16*. slightly raised abaxially, lateral veins inconspicuous, base cune- Thickets along river banks, open grasslands, sandy places near ate, decurrent, margin entire, sparsely 1–3-serrulate, apex acute, seashores; below 500 m. S Guangdong, Hainan, S Taiwan [tropical rarely acuminate, mucronulate. Flowers solitary in leaf axils, Africa, tropical Asia, Australia]. 3. VIOLA Linnaeus, Sp. Pl. 2: 933. 1753.

堇菜属 jin cai shu Herbs perennial or biennial, rarely subshrubs, rhizomatous. Stem developed or absent, sometimes with creeping branches. Leaves simple, alternate or basal, margin entire, dentate, or dissected; stipules small or large, leaflike, free or ± adnate to petioles. Flowers bisexual, zygomorphic, solitary, often dimorphic (cleistogamous flowers later than chasmogamous ones); pedicels axillary, 2-bracteolate. Sepals slightly equal, usually basally auriculate. Petals unequal, anterior petal largest and basally spurred. Filaments free, very short; anthers free or mostly connivent into a sheath around ovary, 2 anterior ones with spurlike or wartlike and nec- tariferous appendages at base, these extending into anterior spur, connectives produced apically into conspicuous, membranous appendages. Ovary 3-carpelled, with many ovuled parietal placentae; styles nearly erect or usually ± curved downward, ± thickened or sometimes gradually tapering toward apex, entire or variously appendaged; style apex and stigma variously shaped. Capsule loculicidally and elastically 3-valved, valves carinate and abaxially thickened. Seeds globose-ovoid, arillate or not, usually smooth; endosperm abundant; embryo straight; cotyledons rather thick, plano-convex. About 550 species: cosmopolitan, chiefly in temperate regions of the N hemisphere; 96 species (35 endemic, three introduced) in China. The identity of Viola curvicalcarata W. Becker & H. Boissieu (Bull. Herb. Boissier, sér. 2, 8: 740. 1908), described from Shaanxi, cannot be ascertained in this treatment because we have not seen any specimens of it. 1a. Anterior petal shallowly saccate but not spurred; stipules free and small; flowers white to violet ...... 5. V. subg. Erpetion 1b. Anterior petal spurred; stipules adnate or free; flowers violet, purple, rose, yellow, or white. 2a. Stipules large, free, leaflike, deeply lobed; anterior petal broadly cuneate, with aerial stem ...... 4. V. subg. Melanium 2b. Stipules small, entire or fimbriate; anterior petal narrowly ovate to linear-oblong, with or without aerial stem. 3a. Flowers purple or white, never yellow; style beaked at apex; stipules adnate or free; plant with or without aerial stem ...... 1. V. subg. Viola 3b. Flowers yellow, never purple; style not beaked at apex; stipules free; plant with aerial stem. 4a. Lateral petals and stigmas beardless; stigmas bilobed; spur short or long ...... 2. V. subg. Dischidium 4b. Lateral petals and stigmas bearded; stigmas capitate; spur very short ...... 3. V. subg. Chamaemelanium

1. Viola subg. Viola 堇菜亚属 jin cai ya shu

Plant with or without aerial stem. Stipules adnate to petioles or free. Flowers violet, purple, pink or white. Anterior petal spurred. Style bearded at apex.

VIOLACEAE 75

About 185 species: mainly in the N hemisphere, occasionally in Oceania; 78 species (27 endemic) in China. 1a. Plants with or rarely without aerial stem in young plants; stipules free (except in V. moupinensis). 2a. Leaves, stipules, and capsules without brown glandular dots; margins of stipules usually entire or remotely denticulate. 3a. Cauline leaves cordate or reniform. 4a. Cauline leaves basally shallowly cordate; stipules brown, margin entire; flowers violet; fruit 10–14 mm ...... 1. V. mirabilis 4b. Cauline leaves basally usually deeply cordate; stipules green, margin entire or dentate; flowers usually white or purplish; fruit 6–8 mm ...... 2. V. arcuata 3b. Cauline leaves ovate to triangular, oblong-triangular, triangular-cordate, or ovate-cordate. 5a. Cauline leaves ovate-lanceolate, base rounded or shallowly cordate ...... 3. V. henryi 5b. Cauline leaves triangular, base shallowly hastate, cordate, or truncate. 6a. Cauline leaves basally hastate, shallowly cordate, or subtruncate; stipules leaflike, 1.5–4 cm ...... 4. V. raddeana 6b. Cauline leaves basally cordate or truncate; stipules less than 1 cm ...... 5. V. triangulifolia 2b. Leaves, stipules, and capsules with brown glandular dots on both sides; margins of stipules usually fimbriate-laciniate or fimbriate-serrate. 7a. Cauline leaves lanceolate. 8a. Cauline leaves lanceolate or ovate-lanceolate, base truncate or rounded; lower and middle stipules subequaling or shorter than petioles, upper ones exceeding petioles ...... 6. V. montana 8b. Cauline leaves lanceolate or broadly lanceolate, base cuneate; lower stipules membranous, elliptic-lanceolate, base amplexicaul, margin usually remotely serrulate, upper ones lanceolate, apex acute, adnate to petiole at base, margin usually pinnatifid ...... 7. V. websteri 7b. Cauline leaves cordate, ovate-cordate, triangular-ovate, or ovate. 9a. Basal leaves absent at anthesis ...... 8. V. acuminata 9b. Basal leaves present at anthesis. 10a. Cauline leaves triangular-ovate, base broadly cuneate or truncate, decurrent to petiole ...... 9. V. stewardiana 10b. Cauline leaves orbicular-cordate, triangular-cordate, ovate-cordate, ovate, or reniform, base ± cordate. 11a. Cauline leaves orbicular-cordate or reniform, apex acute to rounded. 12a. Plant puberulous; auricles of sepals 0.5–1.5 mm; lateral petals bearded ...... 10. V. rupestris 12b. Plant glabrous; auricles of sepals 2–3 mm; lateral petals glabrous ...... 11. V. sacchalinensis 11b. Cauline leaves triangular-ovate, triangular-cordate, or narrowly ovate-cordate, apex acute to acuminate. 13a. Spur 6–8 mm ...... 12. V. grypoceras 13b. Spur 2–3 mm. 14a. Lateral petals beardless; margin of leaf blade crenate ...... 16. V. obtusa 14b. Lateral petals bearded; margin of leaf blade serrulate. 15a. Rhizome short ...... 13. V. kosanensis 15b. Rhizome stout and long. 16a. Rhizome very stout and very long; leaves relatively larger, usually glabrous, base truncate or slightly cordate; capsule large ...... 14. V. pendulicarpa 16b. Rhizome densely nodded; leaves relatively smaller, pubescent, base cordate; capsule very small ...... 15. V. serrula 1b. Plants usually without aerial stem; stipules free or adnate to petioles at base. 17a. Capsules nearly globose, puberulous, rarely glabrous; peduncle prostrate when mature. 18a. Plant without stolons. 19a. Leaf blade broadly ovate or orbicular, to 8 cm; margin of stipules remotely fimbriate-denticulate; flowers blue-purple with white throat ...... 20. V. collina 19b. Leaf blade oblong-ovate, rarely orbicular, to 6 cm; margin of stipules shortly fimbriate; flowers blue to deep purple ...... 21. V. hirta 18b. Plant with stolons. 20a. Stolons underground; flowers fragrant ...... 17. V. odorata 20b. Stolons aerial. 21a. Leaf blade cordate, apex acuminate, margin obtusely dentate; sepals lanceolate, base auricles 2–2.5 mm; lateral petals bearded ...... 18. V. pilosa 21b. Leaf blade broadly ovate or reniform, apex rounded, margin crenate; sepals ovate, base auricles ca. 1 mm; lateral petals usually glabrous ...... 19. V. hondoensis 17b. Capsules oblong to ellipsoid, glabrous; peduncle erect. 22a. Plant with underground stolons.

76 VIOLACEAE

23a. Plant with bulbs ...... 22. V. bulbosa 23b. Plant without bulbs. 24a. Rhizome slender; leaves small, reniform, apex rounded ...... 23. V. epipsiloides 24b. Rhizome slightly stout; leaves large, cordate, apex acuminate ...... 25. V. diamantiaca 22b. Plant without underground stolon. 25a. Plant with aerial stolons. 26a. Spur 5–7 mm ...... 29. V. formosana 26b. Spur less than 2 mm. 27a. Leaves on stolons usually rosette; stipules mostly free; lower petal the shortest, apex acute. 28a. Leaves basally conspicuously decurrent to petioles ...... 26. V. diffusa 28b. Leaves nearly wingless. 29a. Bracteoles linear ...... 28. V. lucens 29b. Bracteoles lanceolate, 5–7 mm, minutely ciliate ...... 31. V. nagasawae 27b. Stolons elongated, with scattered leaves; stipules nearly free; lower petal usually equaling others, apex rounded. 30a. Stipules usually entire; sepals broad ovate, ca. 5 × 3 mm ...... 27. V. grandisepala 30b. Stipules fimbriate; sepals lanceolate, much narrower. 31a. Leaf blade distinctly spinose on margin. 32a. Leaf blade margin conspicuously spinose at apices of teeth ...... 37. V. mucronulifera 32b. Leaf blade margin shortly spinose in between teeth ...... 33. V. kwangtungensis 31b. Leaf blade without spines on margin. 33a. Leaf blade orbicular or nearly orbicular, apex rounded, margin crenate. 34a. Rhizome usually long, nodes elongated; lateral petals bearded ...... 30. V. davidii 34b. Rhizome densely noded; lateral petals glabrous. 35a. Leaf blade adaxially scabrous, sparsely puberulous ...... 34. V. duclouxii 35b. Leaf blade adaxially shining, glabrous ...... 32. V. sikkimensis 33b. Leaf blade cordate, apex acuminate or acute, margin crenate-serrate. 36a. Rhizome short, densely noded; petioles usually longer than leaf blade ...... 36. V. fargesii 36b. Rhizome long and thick, nodes elongated. 37a. Leaves, petioles, and pedicels densely hirsute; sepals puberulous; petiole usually shorter than leaf blade, leaf blade oblong or oblong-ovate, apex acute or acuminate; lateral petals beardless ...... 35. V. yunnanensis 37b. Leaf blade glabrous or sparsely puberulous; petioles, pedicels, and sepals glabrous, or sometimes petioles ± puberulous. 38a. Leaf blade glabrous; sepals ovate, 2–2.5 mm wide ...... 40. V. nuda 38b. Leaf blade ± puberulous; sepals lanceolate, 1.2–1.8 mm wide. 39a. Leaves not glandular; bracteoles 10–13 mm; flowers violet, purplish, or white; lateral petals bearded ...... 38. V. thomsonii 39b. Leaves and capsules dotlike brown glandular; bracteoles 3–8 mm; flowers purple to light purple; lateral petals glabrous ...... 39. V. sumatrana 25b. Plant without aerial stolons (except in V. moupinensis). 40a. Stipules free; rhizome distinctly thick, densely noded, with numerous brown rootlets at nodes. 41a. Leaves all basal, puberulous; stipules small, 5–8 mm ...... 41. V. rossii 41b. Aerial stems usually present, with leaves and fruits at apex; leaves usually glabrous; stipules large, 1–1.8 cm ...... 42. V. moupinensis 40b. Stipules ± adnate to petioles. 42a. Leaf blade multi-lobed, dissected, or incised. 43a. Leaf blade palmately 5-lobed ...... 43. V. dactyloides 43b. Leaf blade dissected, divided, or incised. 44a. Sepals smaller, auricles less than 2 mm; stipules smaller. 45a. Petals pinkish or pinkish white, lower petal apex acute; leaf usually deeply divided ...... 44. V. dissecta 45b. Petals blue violet or white, lower petal apex obtuse; leaf blade margin incised or slightly divided ...... 47. V. forrestiana 44b. Sepals 10–14 mm, auricles 4–6 mm, apex dentate; stipules larger. 46a. Stipules ca. 1/2 adnate to petioles ...... 46. V. albida 46b. Stipules over 1/2 adnate to petioles. 47a. Leaf blade usually pinnatiparted; stipules ca. 1/2 adnate to petioles, the free part ca. 8 mm; petals white, creamy-white or light purplish ...... 45. V. chaerophylloides

VIOLACEAE 77

47b. Leaf blade usually undivided, margin distinctly coarsely serrate; stipules mostly adnate to petioles, the free part 2–5 mm; petals light pink or bluish purple ...... 55. V. magnifica 42b. Leaves simple, margin crenate or serrate. 48a. Roots dark brown or black; leaf base usually decurrent to petiole. 49a. Leaf blade not decurrent to petioles ...... 68. V. lactiflora 49b. Leaf blade distinctly decurrent to petioles. 50a. Roots black; petioles not obvious or very short when young ...... 48. V. gmeliniana 50b. Roots dark brown; petioles usually much longer than blades. 51a. Spur 5–10 mm; flowers violet, rarely white ...... 49. V. mandshurica 51b. Spur shorter than 3 mm; flowers white ...... 50. V. patrinii 48b. Roots yellowish to grayish white; leaf base usually not decurrent to petiole. 52a. Spur more than 4 mm. 53a. Rhizome short, thick, leaves in summer prominently larger than those in spring. 54a. Leaf blade lanceolate, margin shallowly crenate, base distinctly decurrent to petiole ...... 63. V. philippica 54b. Leaf blade ovate, margin serrate or deeply crenate, base not decurrent to petiole. 55a. Stipules densely sericeous ...... 69. V. hirtipes 55b. Stipules glabrous or shortly pubescent. 56a. Lateral petals glabrous or sparsely bearded. 57a. Leaf blade base usually truncate or slightly cordate; lateral petals usually glabrous .... 76. V. prionantha 57b. Leaf blade usually distinctly cordate. 58a. Leaf apex usually acuminate; stipules ca. 1/2 adnate to petioles; petals white or light violet; spur 4–5 mm ...... 66. V. belophylla 58b. Leaf apex usually obtusely acute; stipules ca. 2/3 adnate to petioles; petals light purplish or whitish violet; spur 5–9 mm ...... 77. V. japonica 56b. Lateral petals distinctly bearded. 59a. Petals usually violet or red-purple. 60a. Ovary pubescent ...... 70. V. phalacrocarpa 60b. Ovary glabrous. 61a. Leaves and pedicels usually glabrous ...... 74. V. macroceras 61b. Leaves and pedicels usually pubescent ...... 61. V. tienschiensis 59b. Petals usually white, rarely pinkish white. 62a. Rhizome usually dark brown and very thick ...... 64. V. mongolica 62b. Rhizome yellowish white, slender. 63a. Bracteoles ciliate ...... 56. V. sphaerocarpa 63b. Bracteoles glabrous. 64a. Sepal and stipules and fruit (ca. 6 mm) smaller; leaf abaxially glabrous ...... 67. V. pekinensis 64b. Sepal and stipules and fruit (ca. 10 mm) larger; leaf abaxially pubescent ...... 65. V. hancockii 53b. Rhizome short, thin, leaves in summer nearly the same size as those in spring. 65a. Leaf blade apex rounded. 66a. Leaf blade orbicular to reniform, adaxially distinctly white variegated along veins, abaxially red-purple ...... 52. V. variegata 66b. Leaf blade orbicular to ovate, adaxially green or variegated along veins, abaxially green ...... 53. V. tenuicornis 65b. Leaf blade apex acute. 67a. Plants without underground stolons; leaves usually glabrous, lateral petals usually glabrous ...... 73. V. violacea 67b. Plants usually with underground stolons; leaves pubescent. 68a. Lateral petals glabrous; leaf blade orbicular-ovate to ovate ...... 71. V. selkirkii 68b. Lateral petals sparsely pubescent; leaf blade triangular-ovate to narrowly ovate ...... 72. V. tokubuchiana 52b. Spur less than 3 mm. 69a. Leaf blade orbicular or suborbicular; rhizome very densely noded. 70a. Lateral petals bearded; spur narrow ...... 51. V. striatella 70b. Lateral petals usually glabrous; spur broad ...... 24. V. brachyceras 69b. Leaf blade ovate, oblong, elliptic, triangular or cordate; rhizome relatively sparsely noded. 71a. Leaf blade ovate or cordate. 72a. Lateral petals bearded ...... 75. V. yunnanfuensis 72b. Lateral petals glabrous ...... 62. V. senzanensis 71b. Leaf blade elliptic, oblong or triangular. 73a. Leaf blade base truncate or cordate; stipules brownish.

78 VIOLACEAE

74a. Leaves triangular-lanceolate, base truncate, usually distinctly decurrent to petiole; spur short ...... 59. V. betonicifolia 74b. Leaves triangular-ovate, base cordate or truncate, slightly or not decurrent to petiole; sepal auricles usually distinct, spur slender ...... 57. V. inconspicua 73b. Leaf blade base rounded or cuneate; stipules whitish. 75a. Lateral petals glabrous. 76a. Leaves glabrous, margin entire or sparsely and shallowly crenate ...... 54. V. kunawarensis 76b. Leaves pubescent, margin crenate ...... 78. V. perpusilla 75b. Lateral petals distinctly bearded. 77a. Upper petals bearded ...... 60. V. trichopetala 77b. Upper petals glabrous ...... 58. V. cuspidifolia 1. Viola mirabilis Linnaeus, Sp. Pl. 2: 936. 1753. Viola alata Burgersdijk; V. alata subsp. verecunda (A. Gray) W. Becker; V. amurica W. Becker; V. arcuata f. radi- 奇异堇菜 qi yi jin cai cans (Makino) Nakai; V. arcuata var. verecunda (A. Gray) Na- Viola brachysepala Maximowicz; V. mirabilis var. kai; V. distans Wallich; V. excisa Hance; V. hupeiana W. Beck- brachysepala (Maximowicz) Regel; V. mirabilis var. brevi- er; V. verecunda A. Gray; V. verecunda f. hensoaensis Kudo & calcarata Nakai; V. mirabilis var. glaberrima W. Becker; V. Sasaki; V. verecunda f. radicans Makino; V. verecunda var. mirabilis f. latisepala W. Becker; V. mirabilis var. platysepala semilunaris Maximowicz. Kitagawa; V. mirabilis var. subglabra Ledebour. Herbs perennial. Rhizome procumbent, brown, ca. 2 mm Herbs perennial, acaulescent before anthesis, later grad- in diam., with numerous fibrous roots, producing several stems ually caulescent, 6–23 cm tall. Rhizome erect or oblique, to 4 or stolons. Stems usually several fasciculate, greenish, to 35 cm cm, 3–7 mm in diam., conspicuously noded, much branched in tall; internodes long. Stolons flexible, to 40 cm, producing upper part, with dense brown or red-brown remains of scalelike adventitious roots at nodes; internodes long. Stipules green, stipules; roots numerous, brown, harder when dry. Stem erect, linear, lanceolate, ovate or spatulate, 5–25 × 1–8 mm, glabrous, puberulous or glabrous, with only 1 leaf at middle, densely margin usually entire, or remotely denticulate or ciliate, apex leaved in upper part. Stipules large; basal ones red-brown, scale- acuminate or acute; basal leaves long petiolate, petioles 5–20 like, ovate, ca. 1 × 0.5 cm, apex obtuse or acuminate; upper cm, narrowly winged; petioles of leaves on stem and stolon ones brownish, broadly lanceolate, 0.8–1.7 × 0.3–0.5 cm, gla- shorter; blades of basal leaves deep green, triangular-cordate or brous, margin entire; those of cauline leaves lanceolate, margin ovate-cordate, 1.5–3 × 2–5.5 cm, both surfaces usually glabrous usually ciliate; petioles of basal leaves longer, 5–15 cm, puberu- or sparsely puberulous along veins abaxially, base usually lous or glabrous, narrowly winged; those of cauline leaves un- broadly cordate, rarely deeply cordate, sinus lunate, lobes large equal in length, middle ones ca. 8 cm, upper ones very short or and spreading, margin sparsely and shallowly serrate, apex nearly absent; leaf blade broadly cordate or reniform, 3–5 × 4– acute, rarely acuminate; blades of leaves on stem and stolon 6 cm, abaxially puberulous along veins, adaxially puberulous, similar to those of basal leaves. Flowers light purplish or white, base broadly cordate, margin shallowly crenate, often involute all produced from axils of leaves on stem and stolon; pedicels at anthesis, apex rounded or mucronate. Flowers purplish or vio- long, 2-bracteolate above middle; bracteoles linear. Sepals let, large; those in basal leaf axils usually not fruiting, pedicels ovate-lanceolate, ca. 4 mm, apex acute, basal auricles very to 10 cm, with 2 linear bracteoles in upper part; those in cauline short, suborbicular, margin narrowly membranous. Petals nar- leaf axils fruiting, pedicels short, with bracteoles at or below rowly obovate, ca. 7.5 mm, lateral ones dark purple striate, in- middle. Sepals oblong-lanceolate, ovate-lanceolate, or lanceo- side sparsely bearded or glabrous, anterior one shorter, conspic- late, 7–16 × 2–4 mm, apex acute, basal auricles ca. 2 mm, 3- uously dark purple striate, apex rounded; spur short, ca. 2 mm; veined, margin ciliate or subglabrous, apex obtuse, outer 3 usu- spur of 2 anterior stamens short and stout, subequaling anthers, ally conspicuously longer and broader than inner 2. Petals obo- apex rounded. Ovary glabrous; styles clavate, base slightly vate, lateral ones distinctly bearded, anterior one to 2 cm (spur geniculate, gradually thickened upward; stigmas 2-lobed, lobes included), spur robust, ca. 5 mm, usually curved upward, rarely thick, erect, raised in central part, shortly beaked at base in be- straight, apex obtuse. Ovary glabrous; styles suberect at base tween lobes in front, beak turning upward, with a rounded stig- or slightly curved forward, slightly thickened in upper part, ma hole at tip. Capsule oblong, 6–8 mm, ca. 3 mm in diam., gla- slightly curved and shortly beaked at apex, not papillose; stigma brous, apex acute. Seeds yellowish, ovoid, ca. 1.5 mm, ca. 1 hole narrow. Capsule ellipsoid, 1–1.4 cm, glabrous. Fl. May, fr. mm in diam., with a membranous wing on one side at base. Fl. May–Aug. 2n = 20. Mar–Jun. 2n = 24*. Broad-leaved forests, needle-leaved and broad-leaved mixed Moist and marshy places, grasslands, grassy places on mountain forests, forest margins, thickets, grassy slopes; below 2000 m. Gansu, slopes, thickets, forest margins, fields, alongside houses; usually at low- N Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Ningxia, Shanxi er elevation, below 3000 m. Anhui, Chongqing, Fujian, Gansu, Guang- [Japan, Korea, Mongolia, Russia]. dong, Guangxi, Guizhou, Heilongjiang, Henan, Hubei, Hunan, Jiangsu, 2. Viola arcuata Blume, Bijdr. 58. 1825. Jiangxi, Jilin, Liaoning, Shaanxi, Shandong, Sichuan, Taiwan, Yunnan, Zhejiang [Bhutan, India, Indonesia, Japan, Korea, Malaysia, Mongolia, 如意草 ru yi cao Myanmar, Papua New Guinea, Nepal, Russia, Thailand, Vietnam].

VIOLACEAE 79

In FRPS (51: 105. 1991), the name Viola hamiltoniana D. Don vate, 7–8 mm, with brown glandular dots, margin slightly was misapplied to this species. repand, lateral ones glabrous, anterior one shorter, ca. 7.5 mm 3. Viola henryi H. Boissieu, Bull. Herb. Boissier, sér. 2, 1: (spur included), deep blue-purple striate; spur very short, ca. 2 1075. 1901. mm, apex rounded, slightly exceeding basal auricles of sepals. Anthers ca. 1.8 mm, appendage of connectives ca. 1.2 mm, spur 巫山堇菜 wu shan jin cai of 2 anterior stamens subtriangular, very short, only ca. 1 mm, apex obtuse. Ovary glabrous, with glandular dots; styles slightly Herbs perennial, to 30–40 cm tall. Rhizome obliquely as- slender at base, conspicuously geniculate forward, thicker in cending or erect, to 7 cm, 2–4 mm in diam., with numerous upper part; stigmas 2-lobed, lobes erect, thick, shortly beaked at long, slender rootlets. Stems solitary or several, erect, smooth, base in between 2 lobes, with a conspicuous stigma hole at tip glabrous, leafless at base, few leaved at middle and lower parts, of beak. Capsule oblong, ca. 1 cm. Fl. Jul, fr. Aug–Sep. 2n = densely leaved at apex. Stipules green, ovate, 5–7 mm, margin 24. fimbriate-dentate, apex acuminate; petioles 1–8 cm, those of leaves at middle and lower parts longer, usually slightly longer Thickets and forests along riversides, moist grasslands; below than or subequaling blades, those of leaves at apex conspicu- 1200 m. Heilongjiang, Jilin, Nei Mongol [Japan, Korea, Russia]. ously shorter than blades; blades adaxially green, ovate or This species is very rare and possibly endangered. ovate-lanceolate, 3.5–8 × 2–4 cm, both surfaces sparsely pu- berulous or subglabrous, base shallowly cordate or rounded, 5. Viola triangulifolia W. Becker, Bull. Misc. Inform. Kew slightly decurrent, margin obtusely serrate, apex long acumi- 1929: 202. 1929. nate. Flowers in leaf axils at apex, purplish violet; pedicels 三角叶堇菜 san jiao ye jin cai much shorter than leaves, slender, 2-bracteolate at upper part; bracteoles opposite, subulate. Sepals narrowly linear, ca. 4 × 0.7 Herbs perennial, caulescent, 13–35 cm tall. Rhizome usu- mm, margin narrowly membranous, apex acuminate, basal ally obliquely ascending, deep brown, short, stout, densely auricles very short, apex truncate. Petals oblong-obovate, upper noded, with numerous stout rootlets. Stems solitary or several ones 1–1.2 cm × ca. 4 mm, lateral ones ca. 1.1 cm × 3 mm, gla- fasciculate, erect, slender, glabrous. Basal leaves 2–5, usually brous, anterior one ca. 9 mm (spur included); spur shallowly withered early; petioles 3–6.5 cm; blades broadly ovate or ovate, saccate, 2–3 mm, ca. 2.5 mm in diam. Anthers ca. 2 mm, ap- 1–2 × 1.5–2.8 cm, base cordate, apex acute. Cauline leaves pendage of connectives ca. 2 mm, spur of 2 anterior stamens long petiolate; stipules free, lanceolate or linear-lanceolate, 0.5– shortly angular, ca. 2 mm, ca. 2 mm in diam. Ovary ovoid- 1 cm, herbaceous, margin entire or remotely denticulate, often globose, glabrous; styles clavate, slightly geniculate forward at ciliate; blades ovate-triangular to narrowly triangular, 2–5 × 2– base; stigmas margined on lateral sides and abaxially, shortly 3.5 cm at base, both surfaces glabrous, base cordate or truncate, beaked in front, beak nearly erect, with a rounded stigma hole margin shallowly serrate, apex acute. Flowers solitary in axils at tip. Fl. May, fr. Jul. of cauline leaves, white, purple striate, small; pedicels usually subequaling leaves, sometimes exceeding leaves, slender, 2- ● Shaded and moist places in forest understories of mountain valleys; 1200–1800 m. W Hubei, NW Hunan, E Sichuan. bracteolate at upper part; bracteoles opposite, linear. Sepals ovate-lanceolate or lanceolate, ca. 3 × 0.8–1.2 mm, apex acumi- 4. Viola raddeana Regel, Bull. Soc. Imp. Naturalistes Moscou nate, basal auricles ca. 0.5 mm, margin narrowly membranous. 34(2): 463. 1861. Upper petals narrowly obovate, 5–7.5 × ca. 2.5 mm, lateral ones 立堇菜 li jin cai oblong, 5–8 × ca. 2 mm, bearded, anterior one shorter, spatu- late, ca. 6 mm (spur included); spur shallowly saccate, 1–1.5 Viola deltoidea Yatabe. mm, ca. 2 mm in diam. Anthers ca. 2 mm, appendage of con- nectives ca. 1.5 mm, spur of 2 anterior stamens shorter, square, Herbs perennial, glabrous. Rhizome short, densely noded, subtriangular, ca. 1.1 × 0.8 mm. Ovary ovoid-globose, ca. 1.5 with long, slender roots. Stems usually several fasciculate, mm, glabrous; styles clavate, ca. 1.3 mm, base slightly genicu- rarely solitary, erect or obliquely ascending, ca. 30 cm tall at late forward, conspicuously thickened upward; stigmas conspic- anthesis, elongated after anthesis, to 100 cm tall, slender, gla- uously thickened on lateral and abaxial sides, margin obliquely brous, usually simple or sometimes branched. Basal leaves spreading, flat in central part, shortly beaked in front, with a withered early; stipules of cauline leaves free, leaflike, narrowly larger stigma hole at tip of beak. Capsule ellipsoid, small, 5–6 lanceolate, large, 1.5–4 cm, subequaling or exceeding petioles, mm, glabrous. Fl. Apr–May, fr. May–Sep. glabrous, margin subentire or usually irregularly 1–3-lobed in lower part, apex slightly obtuse or acute; petioles short, 1–3 cm, ● Streamsides in mountain valleys, forest margins, roadsides; narrowly winged in upper part; blades of cauline leaves oblong- 200–1800 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, triangular, 1.5–7 × 1–2 cm, both surfaces glabrous, base nearly Hunan, Jiangxi, Zhejiang. hastate, shallowly cordate, or subtruncate, margin remotely and 6. Viola montana Linnaeus, Sp. Pl. 2: 935. 1753. shallowly crenate or subentire, apex ± acute or obtuse. Flowers bluish purple, small, long pedicellate; pedicels produced from 高堇菜 gao jin cai cauline leaf axils, usually not exceeding leaves, 2-bracteolate at Viola elatior Fries; V. montana var. elatior (Fries) Regel. upper part; bracteoles linear, small. Sepals lanceolate or sub- ulate, 4–4.5 mm, apex slightly acuminate, basal auricles very Herbs perennial. Stem erect, to 50 cm tall, robust, puberu- short, apex truncate and remotely crenate. Petals oblong-obo- lous. Stipules leaflike, lanceolate or ovate-lanceolate, large,

80 VIOLACEAE

lower and middle ones subequaling or shorter than petioles, lous; blade cordate, ovate-cordate, or ovate, 1.5–5.5 × 1.5–4.5 upper ones exceeding petioles, 2–5.5 cm, base often 2- or 3- cm, densely dotlike brown glandular on both sides, sparsely lobed, margin usually entire in upper half, irregularly dentate in puberulous along veins, base usually cordate, rarely truncate, lower half, apex acute; petiole 1.5–4 cm, narrowly winged in margin obtusely serrate and shortly ciliate, apex acute or acu- upper half; leaf blade lanceolate or ovate-lanceolate, 2–7 × 1.5– minate. Flowers whitish violet or white, long pedicellate; pedi- 2.5 cm, both surfaces minutely puberulous, with dense brown cels usually exceeding leaves, slender, puberulous, with 2 linear glandular dots, base truncate or shallowly cordate, slightly de- bracteoles above middle or near flower. Sepals linear-lanceo- current to petiole, margin obtusely dentate or ciliate, apex acute. late, 7–12 × 1.5–2.5 mm, outer 3 longer and broader, apex acu- Flowers purplish violet, whitish at throat, large; pedicels ex- minate, basal auricles 2–3 mm, apex truncate or sometimes 1- ceeding or subequaling leaves, with 2 linear bracteoles above or 2-dentate, puberulous adaxially and along margin, 3-veined. middle. Sepals lanceolate, 1–1.4 cm, apex acuminate, basal Petals dotlike brown glandular, upper petals subequaling lateral auricles short, apex shallowly dentate. Petals obovate, lateral ones, curved upward, usually glabrous, rarely bearded, lateral ones ca. 1.8 cm, distinctly bearded, anterior one shorter, apex ones distinctly bearded, anterior one often purple veined, 0.9– emarginate, ca. 2 cm (spur included); spur greenish, longer than 1.6 cm (spur included); spur usually straight, 1.5–5 mm, sac- basal auricles of sepals; spur of 2 anterior stamens 3–4 mm, cate, apex obtuse; spur of 2 anterior stamens short and obtuse, base thicker, apex thinner. Styles slightly geniculate at base, ca. 1.5 mm. Ovary conic, glabrous; styles slightly curved for- thickened upward and clavate, densely papillose in upper part, ward at base, gradually thickened, with several series of papil- curved forward and uncinately beaked at apex, tip of beak lose protuberances at apex, shortly beaked at apex, tip of beak slightly thickened along margin, with a larger stigma hole at slightly curved upward, with a larger stigma hole. Capsule middle of beak. Capsule glabrous, conspicuously angled, apex ellipsoid, ca. 1 cm, glabrous, usually dotlike yellow-brown mucronate, oblong. Fl. May, fr. Jun–Aug. 2n = 40. glandular, apex acuminate. Fl. May–Jun, fr. Jun–Sep. Forest margins, forests, grasslands on mountain slopes; 1100– Mixed forests, forest margins, thickets, grasslands on mountain 1200 m. Xinjiang [Kazakhstan, Kyrgyzstan, Russia, Tajikistan, Uzbeki- slopes, moist places and shaded and moist grasslands in stream valleys; stan; Europe]. 400–2500 m. Anhui, Gansu, Hebei, Heilongjiang, Henan, Hubei, Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan, 7. Viola websteri Hemsley, J. Linn. Soc., Bot. 23: 56. 1886. Zhejiang [Japan, Korea, Mongolia, Russia]. 蓼叶堇菜 liao ye jin cai 1a. Spur 1.5–3.5 mm; upper petals beardless Herbs perennial. Rhizome robust, with dense white root- ...... 8a. var. acuminata lets. Stem erect, usually simple, 30–40 cm tall, puberulous, 1b. Spur 4.5–5 mm; upper petals bearded ...... 8b. var. pilifera without leaves in lower part but stipulate. Lower stipules 8a. Viola acuminata var. acuminata elliptic-lanceolate, membranous, base amplexicaul, margin usu- ally remotely serrulate; upper ones lanceolate, margin usually 鸡腿堇菜(原变种) ji tui jin cai (yuan bian zhong) pinnatifid, apex acute, adnate to petioles at base; petiole short; Viola acuminata subsp. austroussuriensis W. Becker; V. blade lanceolate or broadly lanceolate, 5–12 × 1.2–5 cm, acuminata var. austroussuriensis (W. Becker) Kitagawa; V. abaxially subglabrous or puberulous along veins, veins reaching acuminata var. brevistipulata (W. Becker) Kitagawa; V. acu- margin, adaxially and along margin sparsely puberulous, base minata var. intermedia Nakai; V. austroussuriensis (W. Becker) cuneate, decurrent to petiole, margin remotely serrate, apex Komarov; V. canina Linnaeus var. acuminata (Ledebour) acute. Flowers solitary in leaf axils, purple, small; pedicels Regel; V. micrantha Turczaninow (1832), not Presl (1822); V. shorter than leaves, with 2 bracteoles at or above middle. Sepals narrowly lanceolate, apex acuminate, basal auricles short, apex micrantha lus. austroussuriensis (W. Becker) W. Becker; V. truncate. Petals subequal, obovate-spatulate, apex rounded, lat- micrantha prol. brevistipulata W. Becker; V. micrantha lus. eral ones bearded, anterior one ca. 1.3 cm (spur included); spur kiautschauensis W. Becker; V. turczaninowii Juzepczuk. straight, short, ca. 2 mm, apex obtuse. Ovary glabrous; styles Stipules leaflike, 1–3.5 cm × 2–8 mm, herbaceous, oblong, clavate, slightly curved forward at base, apex uncinate and lanceolate or oblong, usually pinnatifid and fimbriate or shal- papillose. Capsule oblong, 1.2–1.5 cm, smooth, glabrous, apex lowly divided, ciliate along margin, both surfaces dotlike brown acute. Seeds yellow, narrowly ovoid. Fl. May, fr. Jun–Aug. glandular, sparsely puberulous along veins. Upper petals gla- Sparse mountain forests; 500–900 m. E Jilin [Korea]. brous, spur usually straight, 1.5–3.5 mm, saccate; spur of 2 an- terior stamens short and obtuse, ca. 1.5 mm. Fl. May–Jun, fr. 8. Viola acuminata Ledebour, Fl. Ross. 1: 252. 1842. Jun–Sep. 2n = 20. 鸡腿堇菜 ji tui jin cai Mixed forests, forest margins, grasslands on mountain slopes, moist places in stream valleys; 400–2500 m. Gansu, Hebei, Heilong- Herbs perennial, usually without basal leaves. Rhizome jiang, Henan, Hubei, Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, erect or oblique, robust, with numerous brownish roots. Stem Shandong, Shanxi, Sichuan [Japan, Korea, Mongolia, Russia]. usually 2–4 fasciculate, erect, 10–40 cm tall, glabrous or white puberulous in upper part. Stipules leaflike, 1–3.5 cm × 2–8 mm, 8b. Viola acuminata var. pilifera C. J. Wang, Acta Bot. Yun- herbaceous, usually pinnatifid and fimbriate, or shallowly divi- nan. 13: 257. 1991. ded, ciliate along margin, both surfaces dotlike brown glandu- 毛花鸡腿堇菜 mao hua ji tui jin cai lar, sparsely puberulous along veins; lower petioles to 6 cm, upper ones shorter, 1.5–2.5 cm, glabrous or sparsely puberu- Stipules lanceolate, remotely denticulate, puberulous.

VIOLACEAE 81

Flowers larger, spur long and thick, 4.5–5 × 2–3 mm. Upper ones much longer than blades, upper ones ca. as long as blades; petals inside bearded, spur of 2 anterior stamens long angular, blade orbicular or ovate-orbicular, 0.8–1.7 × 0.7–1.8 cm, both 4–4.5 mm. Fl. and fr. May–Sep. surfaces puberulous, with brown glandular dots when dry, base ● Forest margins, thickets, shaded and moist grasslands in stream shallowly cordate, margin regularly crenate, apex obtuse. Flow- valleys; 1600–1800 m. Gansu. ers solitary in upper leaf axils purple or purplish; pedicels much exceeding leaves, 2.5–6 cm, slender, sparsely puberulous, 2- 9. Viola stewardiana W. Becker, Repert. Spec. Nov. Regni bracteolate above middle; bracteoles subulate or linear, 4–6 Veg. 21: 237. 1925. mm, fimbriate-dentate at middle and lower parts. Sepals 庐山堇菜 lu shan jin cai broadly lanceolate or lanceolate, 4–7.5 × 1–2.5 mm, 3-veined, with dense brown glandular dots, apex acute, basal auricles Herbs perennial; tap roots long. Rhizome robust, densely square, shorter, 0.5–1.5 mm, with 2 or 3 obtuse teeth at apex. noded. Underground part of stem procumbent, strongly ligni- Petals narrowly oblong-ovate, veins deeply colored, lateral fied, very hard, often producing new plants; stems usually sev- petals 0.8–1 × ca. 4 mm, distinctly bearded, upper ones sub- eral fasciculate, obliquely ascending, 10–15 cm tall, longitu- equaling lateral ones, anterior one shorter, 1.3–1.4 cm (spur dinally angled, glabrous. Basal leaves rosulate; petiole to 5.5 included), with dense brown glandular dots; spur cylindric, 3–4 cm; blade triangular-ovate, 1.5–3 × 1.5–2.5 cm, both surfaces mm, ca. 2 mm in diam., apex rounded, horizontally straight or with minute brown glandular dots, veins conspicuously raised slightly curved upward. Anthers ca. 1.3 mm, connective ap- abaxially, base broadly cuneate or truncate, decurrent to petiole, pendages ca. 0.7 mm; 2 anterior stamens ca. 3 mm, spur margin crenate, glandular at apices of teeth, apex mucronate. broadly triangular, ca. 2.5 × 0.8 mm. Ovary deep brown, ovoid, Stipules of cauline leaves brown, lanceolate or linear-lanceo- styles clavate, evenly thick, slightly narrower only at base, late, basal ones 1–1.2 cm, upper ones only ca. 0.5 cm, margin slightly geniculate forward at base, curved forward at tip and long fimbriate, apex long acuminate; lower petioles subequal- beaked, conspicuously papillose abaxially and on lateral sides ing blades, upper ones shorter than blades, narrowly winged; of beak; beak horizontal, mouth slightly curved upward, with a blades narrowly ovate, rhombic, or triangular-ovate, ca. 4.5 × larger stigma hole. Capsule oblong, 5–7 mm, 3–4 mm in diam., 2–3 cm, base cuneate, apex mucronate or acuminate. Flowers in with remains of styles at apex, puberulous or glabrous. upper leaf axils, light violet, long pedicellate; pedicels sub- equaling or slightly exceeding leaves, with 2 linear bracteoles Forests, grassy slopes, meadows in river valleys, sandy grass- slightly above middle. Sepals narrowly ovate or oblong-lan- lands; 1000–2500(–4000) m. SE Gansu, S Shaanxi, S Shanxi, Xinjiang [Kashmir, Kazakhstan, Kyrgyzstan, Mongolia, N Pakistan, Russia, ceolate, 3–3.5 mm, apex mucronate, basal auricles short, gla- Tajikistan; SW Asia (Caucasus), Europe]. brous, margin entire, apex rounded. Petals conspicuously emar- ginate at apex, upper ones spatulate, ca. 8 mm, lateral ones 1a. Plants smaller; stipules usually ovate or oblong, glabrous, anterior one narrowly obovate, ca. 1.4 cm broadly lanceolate, 3–5 × 1.5–2.5 mm, (spur included); spur ca. 6 mm, curved downward, apex obtuse; margin fimbriate-serrate or sparsely 2 anterior stamens spurless. Ovary ovoid-globose, glabrous; emarginate-serrate ...... 10a. subsp. rupestris styles slightly geniculate forward at base, gradually thickened 1b. Plants larger; stipules usually narrowly upward, without appendages at top, shortly beaked, beaks long, linear or subulate, slightly curved upward, with a larger stigma hole at tip. Capsule 5–7 × 0.8–1.3 mm ...... 10b. subsp. licentii subglobose, with scattered brown glands, ca. 6 mm, apex 10a. Viola rupestris subsp. rupestris mucronate. Fl. Apr–May, fr. May–Sep. 2n = 20*. ● Streamsides in mountain valleys, rock crevices; 400–1500 m. 石生堇菜(原亚种) shi sheng jin cai (yuan ya zhong) Anhui, Fujian, SE Gansu, Guangdong, Guizhou, Hubei, Hunan, Jiang- Viola canina Linnaeus var. rupestris (F. W. Schmidt) Regel. su, Jiangxi, S Shaanxi, Sichuan, Zhejiang. Plants 5–10 cm tall; blade orbicular or ovate-orbicular, 10. Viola rupestris F. W. Schmidt, Neuere Abh. Königl. Böhm. 0.8–1.7 × 0.7–1.8 cm, both surfaces puberulous, with brown Ges. Wiss. 1: 60. 1791. glandular dots when dry, base shallowly cordate, margin reg- 石生堇菜 shi sheng jin cai ularly crenate, apex obtuse; stipules of basal leaves usually Herbs perennial, caulescent, 5–10 cm tall, puberulous sparsely arranged, lanceolate, fimbriate-dentate, those of cau- throughout, rarely sparsely puberulous or glabrous. Rhizome line leaves free, ovate or broadly lanceolate, 3–5 × 1.5–2.5 mm, erect, deep brown, short, robust, with nodes and scales; tap root margin fimbriate-serrate or sparsely emarginate-serrate, teeth robust, long; roots brown, much branched in lower part; rootlets with brown glandular dots at apices, apex acuminate. Fl. May– long, slender. Stems many, fasciculate, obliquely ascending or Jun, fr. Jul–Aug. 2n = 20. erect, longitudinally angled. Basal leaves rosulate, long petio- Mountain forests, grassy slopes, meadows in river valleys; 1300– late; cauline leaves similar to basal ones; stipules of basal 2500(–4000) m. Xinjiang [Kashmir, Kazakhstan, Kyrgyzstan, Mon- leaves usually sparsely arranged, lanceolate, fimbriate-dentate, golia, N Pakistan, Russia, Tajikistan; SW Asia (Caucasus), Europe]. those of cauline leaves free, ovate to lanceolate, 3–7 × 0.8–2.5 10b. Viola rupestris subsp. licentii W. Becker, Bull. Misc. mm, margin fimbriate-serrate or sparsely emarginate-serrate, Inform. Kew 1928: 248. 1928. teeth with brown glandular dots at apices, apex acuminate; peti- ole slender, weak, narrowly winged only in upper part, middle 长托叶石生堇菜 chang tuo ye shi sheng jin cai

82 VIOLACEAE

Plant 10–20 cm tall; blades usually wider, subreniform, 11b. Viola sacchalinensis var. alpicola P. Y. Fu & Y. C. Teng, 0.8–1.4 × 1.2–1.8 cm; stipules narrowly long, linear or subulate, Fl. Pl. Herb. Chin. Bor.-Orient. 6: 291. 1977. 5–7 × 0.8–1.3 mm, margin fimbriate-serrate. Fl. Apr–May, fr. 长白山堇菜 chang bai shan jin cai Jul. Viola koraiensis Nakai, Bot. Mag. (Tokyo) 30: 281. 1916. ● Sandy grasslands, forests; 1000–2200 m. SE Gansu, S Shaanxi, S Shanxi. Rhizome densely covered with dead black-brown leaf stip- 11. Viola sacchalinensis H. Boissieu, Bull. Soc. Bot. France ules and petioles. Stem usually short, glabrous. Stipules green, lanceolate, margin serrate or mostly fimbriate-serrate. Petals 57: 188. 1910. light violet or deep violet, broadly oblong-obovate, apex obtuse; 库叶堇菜 ku ye jin cai lateral petals glabrous; spur 2–2.5 mm. Stigma glabrous. Fl. May–Jul, fr. Jul–Sep. Herbs perennial, acaulescent at first, 2–5 cm tall, later gradually caulescent, to more than 20 cm tall, glabrous. Rhi- Alpine grasslands, sparse deciduous forests; 1100–2400 m. S Jilin zome slender, with brown remains of scalelike stipules. Stipules (Changbai Shan) [N Korea]. ovate-lanceolate or narrowly ovate, 0.8–1 cm × 2–4 mm, apex 12. Viola grypoceras A. Gray in Perry, Narr. Exped. China acuminate, adnate to petioles at base inside, margin densely Japan 2: Append. 308. 1856. fimbriate-denticulate; petiole slender, lower ones ca. 4.5 cm, 紫花堇菜 upper ones shorter; blade cordate, ovate-cordate, or reniform, zi hua jin cai 1–2.5 × 1–2.5 cm, both surfaces with brown glandular dots, Viola grayi Franchet & Savatier var. candida H. Boissieu; glabrous or subglabrous, base cordate or broadly cordate, mar- V. grypoceras var. barbata W. Becker; V. grypoceras var. gin obtusely serrate, apex obtuse. Flowers bluish violet, long pubescens Nakai; V. leveillei H. Boissieu; V. sylvestris Lamarck pedicellate; pedicels exceeding leaves, to 5.5 cm, with 2 oppo- var. candida (H. Boissieu) H. Léveillé; V. sylvestris var. site bracteoles near flower. Sepals lanceolate, ca. 5 mm, apex grypoceras (A. Gray) Maximowicz. acuminate, basal auricles 2–3 mm, glabrous, margin entire, apex slightly dentate. Upper and lateral petals obovate, throat white, Herbs perennial, with developed tap root. Rhizome erect, lateral ones distinctly bearded, anterior one oblong, 1.3–1.6 cm brown, short, robust, densely noded. Stems several, erect, (spur included), distinctly veined in throat; spur shorter, ca. 3 obliquely ascending, or nearly procument or creeping, 5–20 cm mm, horizontally straight or slightly curved upward. Ovary tall at anthesis, to 30 cm tall at fruiting, glabrous or puberulous. glabrous, often with glandular dots; styles slightly geniculate Stipules brown, narrowly lanceolate, 1–1.5 cm × 1–2 mm, forward at base, gradually thickened upward, clavate, papillose; margin fimbriate-dentate, teeth 2–5 mm, apex acuminate; peti- oles of basal leaves to 8 cm, those of cauline leaves shorter, beak hooked, with a larger stigma hole curved upward. Capsule puberulous; blades of basal leaves cordate or broadly cordate, ellipsoid, glabrous, apex acute. Fl. Apr–Jul, fr. Jun–Sep. 1–4 × 1–3.5 cm, both surfaces glabrous or subglabrous, rarely Forests, sparse deciduous forests, forest margins, alpine grass- puberulous abaxially, with dense brown glandular dots, base lands; 400–2400 m. Heilongjiang, Jilin, Nei Mongol [Japan, Korea, narrowly sinuate, margin obtusely serrate, apex obtuse or ± Mongolia, Russia]. acute; blades of cauline leaves triangular-cordate or narrowly 1a. Petals oblong-ovate, lateral ones ovate-cordate, 1–6 cm, base shallowly sinuate or broadly tri- distinctly bearded, stigma papillose angular. Flowers purplish to white, fragrant; pedicels produced ...... 11a. var. sacchalinensis from base of stem or cauline leaf axils, 6–11 cm, much ex- 1b. Petals broadly oblong-obovate, lateral ceeding leaves, with 2 linear bracteoles above middle. Sepals ones glabrous, stigma not papillose ...... 11b. var. alpicola lanceolate, ca. 7 mm, with brown glandular dots, apex acute, basal auricles ca. 2 mm, apex truncate, shallowly dentate. Petals 11a. Viola sacchalinensis var. sacchalinensis obovate-oblong, with brown glandular dots, margin repand, lat- eral petals glabrous, anterior one 1.5–2 cm (spur included); spur 库叶堇菜 原变种 ( ) ku ye jin cai (yuan bian zhong) 6–8 mm, ca. 2 mm in diam., usually curved downward, rarely Viola komarovii W. Becker; V. mariae W. Becker; V. straight; 2 anterior stamens long spurred, spur suberect. Ovary miranda W. Becker. glabrous; styles slightly geniculate at base, gradually thickened toward top and clavate; stigmas not papillose, curved forward, Rhizome usually not covered by dead leaf stipules. Stem forming a short beak, with a broad stigma hole at tip of beak. usually longer, to 20 cm tall. Stipules ovate-lanceolate or nar- Capsule ellipsoid, ca. 1 cm, with dense brown glandular dots, rowly ovate, margin densely fimbriate-denticulate. Petals bluish apex acuminate. Fl. Mar–May, fr. May–Aug. 2n = 20*. violet, oblong-ovate, apex obtuse; lateral petals distinctly beard- Grassy slopes, thickets; below 2400 m. Anhui, Fujian, SE Gansu, ed; spur 2.5–3 cm. Stigma papillose. Fl. Apr–Jun, fr. Jun–Sep. Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi, 2n = 20. S Shaanxi, E Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Korea]. Forests, forest margins; 400–2000 m. Heilongjiang, Jilin, Nei In FRPS (51: 31. 1991), the name Viola faurieana W. Becker was Mongol [Japan, Korea, Mongolia, Russia]. misapplied to this species. In FRPS (51: 24. 1991), the name Viola mauritii Teplouchow was One of us (Ohba) prefers to treat plants with stems nearly pro- misapplied to this taxon. cumbent or creeping as Viola grypoceras var. exilis (Miquel) Nakai.

VIOLACEAE 83

13. Viola kosanensis Hayata, Icon. Pl. Formosan. 3: 28. 1913. slightly abruptly acute. Petals obovate-oblong; lateral ones bearded; anterior one distinctly purplish striate in throat; spur 福建堇菜 fu jian jin cai saccate, ca. 2.5 mm, apex rounded. Anthers ca. 2 mm. Ovary Viola fukienensis W. Becker; V. kiangsiensis W. Becker; glabrous, with brown glandular dots; styles clavate, ca. 1.5 mm, V. nagamiana T. Hashimoto; V. shinchikuensis Yamamoto. slightly slender at base and slightly geniculate; stigmas flat at top, slightly thick and elevated behind, with few papillose pro- Herbs perennial, caulescent or acaulescent, 5–15 cm tall. turbances, shortly beaked in front, with larger stigmatic hole at Rhizome erect, yellow-brown, very long, to 12 cm, stout, 4–7 tip of beak, hole spatulate, opening upward. Capsule nearly ob- mm in diam., densely noded, with dense brown remains of long, 4–9 mm, with sparse brown glandular dots. Seeds dark stipules, often branched at apex, with numerous long, slender brown, ovoid-globose, ca. 2 mm, ca. 1 mm in diam. Fl. Mar– fibrous roots. Basal leaves numerous, fasciculate, rosulate; May, fr. May–Sep. stipules free, brown, linear-lanceolate, margin fimbriate-serrate; petiole 1.5–4 cm, very narrowly winged in upper part; blade ● Mountain forests, forest margins, riversides; 300–2400(–3500) ovate or triangular-ovate, 1.3–3.5 × 1–3 cm, both surfaces gla- m. W Hubei, S Shaanxi, Sichuan, NW Yunnan. brous to subglabrous or appressed puberulous, with dense 15. Viola serrula W. Becker, Bull. Misc. Inform. Kew 1928: brown glandular dots, veins conspicuously raised abaxially, 247. 1928. base shallowly cordate or cuneate, margin serrulate, apex ± acute. Flowers deep purple to light purple or white, small, ca. 1 小齿堇菜 xiao chi jin cai cm; pedicels linear, 2–7 cm, slender, soft, glabrous or puberu- Herbs perennial. Stem procumbent or obliquely ascending, lous, 2-bracteolate above middle; bracteoles linear-subulate, 3– 5–15 cm tall. Rhizome erect, hard, densely covered with dense 3.5 mm, margin fimbriate at base. Sepals linear-lanceolate, 5–6 remains of stipules and petioles. Stipules free, brown, narrowly mm, 3-veined, margin entire, apex ± acute, basal auricles ca. lanceolate, margin fimbriate; petiole 2–4 cm; leaf blade cordate 1.5 mm, apex rounded or inconspicuously crenate. Petals ob- or broadly ovate, 1.5–2 × ca. 1 cm, abaxially glabrous, adaxi- long-obovate, lateral ones bearded, anterior one shorter; spur 2– ally pubescent, base widely cordate, margin shallowly crenu- 3 mm. Ovary glabrous; styles slightly geniculate at base; stig- late, apex abruptly acute. Flowers violet or whitish, small; pedi- mas raised, shortly beaked in front, slightly margined abaxially cels long, much exceeding leaves, slender, glabrous, with 2 and adaxially. Capsule oblong, 6–7 mm, glabrous, with brown opposite bracteoles in upper part. Sepals ovate-lanceolate, apex dots. Seeds brown when mature, ovoid-globose, ca. 2 mm, ca. 1 abruptly acute, basal auricles narrower, with brown glandular mm in diam. Fl. Mar–Apr, fr. Aug–Oct. dots. Lateral petals papillose-hairy at base inside; anterior petal ● Forest margins on mountains, shaded and moist places along thickly spurred; spur 2–2.5 mm, horizontally straight. Styles riversides; 200–2700 m. Anhui, Fujian, Guangdong, Guangxi, NE Gui- slender, not geniculate at base, papillose-hairy at apex, with a zhou, Hubei, Hunan, Jiangxi, S Shaanxi, Sichuan, Taiwan, NW Yun- slightly curved short beak in front. Capsule ca. 4 mm, glabrous, nan. with brown glandular dots. Fl. Mar–Apr, fr. Jun–Jul. 14. Viola pendulicarpa W. Becker, Beih. Bot. Centralbl., Abt. ● Rock crevices in sparse forests; 300–2000 m. Chongqing, Gui- 2, 36: 55. 1918. zhou, NE Yunnan.

悬果堇菜 xuan guo jin cai 16. Viola obtusa (Makino) Makino var. tsuifengensis T. Hashi- moto, Acta Tsukuba Bot. Gard. 6: 3. 1987 Viola kansuensis W. Becker; V. kansuensis var. oblonga W. Becker; V. polymorpha C. C. Chang; V. pseudoarcuata C. 翠峰堇菜 cui feng jin cai C. Chang; V. weixiensis C. J. Wang. Herbs perennial, puberulous or rarely glabrescent. Rhi- Herbs perennial, caulescent or sometimes nearly acaules- zome erect or subhorizontal, rather stout with dry brown scales, cent, ca. 15 cm tall, glabrous throughout. Rhizome erect, very 0.5–2 cm, with lateral roots. Stems erect or ascending, 2–12 cm long, to 12 cm, stout, 4–7 mm thick, internodes shortened, at anthesis, usually with 1 or 2 stipulelike scales in between nodes dense and with remains of brown stipules, often branched base and first cauline leaf. Basal leaves few or ephemeral at at top and with numerous long, slender, fibrous roots. Leaves anthesis; stipules brownish, ovate, 8–10 mm, margin fimbriate- numerous, mostly basal, rosulate; stipules free, brown when laciniate; petiole 1.7–6 cm; leaf blade cordate-ovate or rarely dry, linear-lanceolate or subulate, 1.2–2 cm, margin densely, cordate-reniform, 17–30 × 1.6–2.4 cm, margin crenate, apex long fimbriate, with inconspicuous glandular dots on both sides; acute or rarely obtuse. Cauline leaves usually tufted above 1/3 petiole long, to 12 cm, slender, glabrous; leaf blade triangular- of plant; leaf blade mostly triangular-ovate, longer than wide, ovate, ovate, or ovate-cordate, 2–3.5 × 1.6–3 cm, both surfaces base cordate, apex acute or rarely obtuse. Pedicels slender, usu- glabrous but with dense brown glandular dots, base truncate or ally arising from axils of cauline leaves, 2–8 cm, with opposite shallowly cordate, margin serrate, apex abruptly acute or slight- bracteoles near apex; bracteoles linear-triangular. Sepals green ly acuminate. Flowers white or purplish, ca. 1 cm; pedicels with purple tints, lanceolate, 5–7 × 1–1.5 mm, margin mem- filiform, long, 4–7 cm, not or slightly exceeding leaves, slender, branous, apex acuminate; appendages of lateral sepals obtri- 2-bracteolate above middle; bracteoles alternate, linear. Sepals angular, ca. 1.5 mm, margin irregularly dentate. Petals obovate, linear-lanceolate, ca. 6 mm, conspicuously 3-veined, margin to 12 × ca. 5 mm, margin undulate, lateral ones glabrous, an- narrowly membranous, apex acuminate; basal auricles of lower terior petal not recurved; spur 2.5–3 mm. Ovary glabrous; style sepals ca. 1.5 mm, apex rounded; lateral sepals narrower, apex geniculate at base, beaked in front, with a stigma hole at tip of

84 VIOLACEAE

hole, ca. 3 mm, glabrous. Seeds ovoid, ca. 1.2 mm. Fl. Mar– inside deep-colored veined; spur saccate, 2–2.5 mm, usually Apr, fr. May–Sep. sparsely puberulous; spur of 2 anterior stamens ca. 1.5 mm, ● Taiwan (Nantou). angular. Ovary usually puberulous; styles clavate, base slightly geniculate, gradually thickened upward; stigmas ± flat, not mar- Viola obtusa var. obtusa occurs in Japan. gined, very inconspicuously short beaked in front, with smaller 17. Viola odorata Linnaeus, Sp. Pl. 2: 934. 1753. stigma hole at tip of beak. Capsule subglobose, 5–10 mm, puber- ulous or glabrous. Seeds with dotted protuberances on surface, 香堇菜 xiang jin cai conspicuously appendaged on lower lateral side. Fl. Feb–Apr, Herbs perennial, acaulescent, stoloniferous, 3–15 cm tall. fr. May–Sep. 2n = 20. Rhizome erect or oblique, brownish, robust, densely noded, Mountain forests, grasslands, roadsides; 800–3000 m. W Guang- with numerous roots and slender, long stolons produced from xi, Guizhou, Sichuan, SE Xizang, Yunnan [Afghanistan, Bhutan, In- nodes. Leaves basal; leaf blade orbicular or reniform to broadly dia, Indonesia, Kashmir, Malaysia, Myanmar, Nepal, Sri Lanka, Thai- ovate-cordate, smaller at anthesis, 1.5–2.5 × 1.5–2.5 cm, gradu- land]. ally accrescent, to 4.5 × 4.5 cm, both surfaces sparsely puberu- In FRPS (51: 90. 1991), the name Viola canescens Wallich was lous or subglabrous, base deeply cordate, margin crenate, apex misapplied to this species. rounded or ± acute. Flowers deep purple, large, fragrant; pedi- 19. Viola hondoensis W. Becker & H. Boissieu, Bull. Herb. cels long, slender, puberulous or subglabrous, 2-bracteolate at Boissier, sér. 2, 8: 739. 1908. or above middle. Sepals oblong or oblong-ovate, base obtuse or 日本球果堇菜 shallowly dentate, apex obtuse, basal auricles 2–3 mm. Petals ri ben qiu guo jin cai crenate, upper ones obovate, lateral ones inside shortly bearded, Viola hirta Linnaeus var. japonica Maximowicz. anterior one broadly obovate, 1.5–2 cm (spur included); spur 2– Plants 3–8 cm tall at anthesis, pubescent. Rhizome thick, 4 mm, straight or slightly curved; spurs of 2 anterior stamens creeping, nodes closely spaced. Stolons creeping, producing robust, ca. 4 mm. Ovary puberulous; styles slender and straight new individuals. Radical leaves in summer larger than those in at base, thickened upward and slightly compressed, apically spring, usually present in winter; leaf blade to 6–7 × 6–7 cm. curved and with a hooked beak about as long as diameter of Petiole 2–8 cm, pubescent. Stipules broadly lanceolate, margins styles, with a smaller stigma hole at tip of beak. Capsule glo- fimbriate; leaf blade orbicular-cordate to reniform, 1.5–4 × 1.5– bose, densely puberulous. Fl. Apr–May, fr. Jun–Sep. 2n = 20, 4 cm, both surfaces pubescent, base cordate, margin crenate, 60. apex rounded. Pedicel radical or cauline, 3–8 cm. Flowers light Cultivated in large cities including Beijing, Guangzhou, Shanghai, violet or white, 1–1.5 cm in diam. Sepals ovate, pubescent, and Tianjin. Guangdong, Hebei, Shaanxi, Zhejiang [native to N Africa, apex obtuse, basal auricles ca. 1 mm. Petals 1–1.3 cm; lateral C and SW Asia, and Europe]. ones sparsely shortly bearded or glabrous, slightly purple stri- ate; lowest petal purple-striate; spur 3–4.5 mm, 2–3 mm longer 18. Viola pilosa Blume, Catalogus, 57. 1823. than base auricles. Ovary pilose; style hooked at apex. Pedicel 匍匐堇菜 pu fu jin cai curved in fruit. Capsule globose, 5–6 mm in diam., densely pubescent. Fl. Mar–May, fr. Jul–Sep. 2n = 20. Viola pogonantha W. W. Smith; V. serpens Wallich ex Gingins; V. serpens subsp. gurhwalensis W. Becker; V. serpens Mixed forests near streams; 900–1300 m. Chongqing, Hubei, Hunan, Jiangxi, S Shaanxi, Zhejiang [Japan, Korea]. var. pseudoscotophylla H. Boissieu. 20. Viola collina Besser, Cat. Hort. Cremenecr. 151. 1816. Herbs perennial, acaulescent or with very short stem. Rhi- zome erect or obliquely ascending, 3–5 cm × 1.5–4 mm; inter- 球果堇菜 qiu guo jin cai nodes conspicuous. Stolon elongated, slender, glabrous, with Herbs perennial, 4–9 cm tall at anthesis, to 20 cm at fruit- evenly scattered leaves. Leaves nearly basal; stipules mostly ing. Rhizome erect or oblique, yellow-brown, 2–6 cm, robust, free, brown or green, lanceolate, margin long or shortly fim- thick, densely noded, apex often branched; roots numerous, briate-dentate, apex long acuminate; petioles nearly as long as brownish. Leaves basal, rosulate; stipules lanceolate, 1–1.5 cm, blades or lower ones much longer than blades, densely ap- membranous, base adnate to petioles, margin remotely fim- pressed hirsute, more densely hairy in upper part; leaf blade briate-denticulate, apex acuminate; petiole narrowly winged, 2– greenish on both surfaces, ovate or narrowly ovate, 2–6 × 1–3 5 cm at anthesis, to 19 cm at fruiting, appressed puberulous or cm, sparsely white stiffly hairy, more densely so along veins rarely glabrescent; leaf blade broadly ovate or suborbicular, 1– abaxially, base narrowly and deeply sinuate, lateral auricles con- 3.5 × 1–3 cm, conspicuously accrescent at fruiting, to 8 × ca. 6 spicuous, margin densely and shallowly obtusely dentate, apex cm, both surfaces sparsely puberulous or rarely glabrescent, caudate-acuminate or acute. Flowers purplish or white, medium- base shallowly or deeply and narrowly sinuate, later cordate, sized; pedicels usually exceeding leaves, sparsely puberulous or margin shallowly and obtusely serrate, apex obtuse or acute, subglabrous, 2-bracteolate above middle; bracteoles linear. Se- rarely acuminate. Flowers purplish, ca. 1.4 cm, long pedicellate; pals lanceolate, 6–7.5 × ca. 2.5 mm, apex acute, basal auricles pedicels with 2 bracteoles ca. 6 mm at or below middle, pu- 2–2.5 mm, outside usually sparsely white puberulous, margin berulous or rarely glabrescent. Sepals oblong-lanceolate or lan- ciliate, apex remotely and shallowly dentate. Petals oblong-obo- ceolate, 5–6 mm, ciliate and glandular, basal auricles short and vate, base narrower, lateral ones bearded, anterior one shorter, obtuse. Petals whitish at base, upper and lateral ones obtuse at

VIOLACEAE 85

apex, lateral ones bearded or subglabrous, spur of anterior one shortly fimbriate and glandular; petiole 3–7 cm, densely white, shorter, ca. 3.5 mm, slightly curved upward, apex obtuse. puberulous; leaf blade smaller at anthesis, ovate or ovate- Ovary puberulous; styles geniculate at base, gradually thick- cordate, 1.5–2.5 × 1–2 cm, both surfaces puberulous, base shal- ened upward, often sparsely papillose, apically curved down- lowly cordate to deeply cordate, margin obtusely serrate, apex ward forming a hooked beak, with a smaller stigma hole at tip rounded or ± acute. Petiole after anthesis to 10 cm; leaf blade of beak. Capsule globose, densely white puberulous or rarely accrescent after anthesis, oblong-ovate or suborbicular, to 6 cm, glabrescent; fruit stalk usually curved downward when mature. base deeply cordate. Flowers deep purple, large, not fragrant; pedicels usually longer than blades, puberulous, with 2 linear Forests, forest margins, thickets, grasslands, grassy slopes, moun- bracteoles below middle. Sepals oblong, apex obtuse, basal tain valleys, shaded and moist places; below 2800 m. Anhui, Chong- qing, Gansu, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Jiangsu, auricles short, base rounded, margin sparsely fimbriate and Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi, Si- glandular. Petals oblong-obovate, lateral ones bearded, anterior chuan, NW Yunnan, Zhejiang [Japan, Korea, Mongolia, Russia, Tajik- one emarginate at apex, 1.5–1.7 cm (spur included); spur red- istan; Europe]. purple, ca. 5 mm, apex obtuse, usually curved upward; spurs of 2 anterior stamens ca. 4 mm, slender. Ovary globose, puberu- 1a. Both leaves and capsules puberulous ...... 20a. var. collina lous; styles slender and slightly curved forward at base, thick- 1b. Either leaves or capsules glabrous. ened upward, apically curved and with a hooked beak, with a 2a. Petioles and leaf blades glabrous; smaller stigma hole at tip of beak. Capsule globose, puberulous. capsules puberulous ...... 20b. var. intramongolica Fl. Apr–May. 2n = 20. 2b. Petioles and leaf blades puberulous; capsules glabrous Forest margins, grasslands, thickets; 1100–1700 m. Xinjiang ...... 20c. var. glabricarpa [Russia; Europe]. 20a. Viola collina var. collina 22. Viola bulbosa Maximowicz, Mélanges Biol. Bull. Phys.- Math. Acad. Imp. Sci. Saint-Pétersbourg 9: 748. 1876. 球果堇菜(原变种) qiu guo jin cai (yuan bian zhong) 鳞茎堇菜 Viola hirta Linnaeus var. collina (Besser) Regel; V. micro- lin jing jin cai donta Chang. Viola bulbosa var. franchetii H. Boissieu; V. bulbosa Petioles, leaf blades, pedicels, and capsules puberulous. subsp. tuberifera (Franchet) W. Becker; V. filifera Komarov; V. Fl. Mar–May, fr. Jun–Aug. 2n = 20. hookeri Franchet (1886), not Thomson & W. Becker (1916); V. multistolonifera C. J. Wang; V. tuberifera Franchet; V. tuberi- Forests, forest margins, thickets, grassy slopes, mountain valleys, fera var. brevipedicellata S. Y. Chen; V. tuberifera var. pseudo- moist places along roadsides; below 2800 m. Anhui, Chongqing, Gansu, palustris H. Léveillé. Guizhou, Hebei, Heilongjiang, Henan, Hubei, Jiangsu, Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan, NW Yun- Herbs perennial, dwarf, shortly caulescent, 2.5–4.5 cm tall. nan, Zhejiang [Japan, Korea, Mongolia, Russia, Tajikistan; Europe]. Rhizome erect, long, slender, with numerous underground 20b. Viola collina var. intramongolica C. J. Wang, Acta Bot. stolons and a small bulb in lower part; bulb 5–6 mm in diam., Yunnan. 13: 257. 1991. consisting of 4–6, white, fleshy boat-shaped scales, with numer- ous fibrous roots in lower part. Leaves clustered at apex of 光叶球果堇菜 guang ye qiu guo jin cai stem; stipules 1–1.4 cm, only 1/3 adnate to petioles, free part Petioles and leaf blades glabrous. Sepals oblong, both sur- linear-lanceolate, ciliate or glabrous, apex acute; petiole nar- faces glabrous, with yellow-brown, small glands only along rowly winged, usually shorter than or subequaling blade, pu- margin proximally, basal auricles sparsely ciliate. Capsules pu- berulous; leaf blade oblong-ovate, suborbicular or reniform, 1– berulous. Fr. Aug. 2.5 cm × 5–14 mm, glabrous, or white puberulous abaxially particularly on young leaves, base cuneate or narrowly cordate, ● Larix forests, grasslands; 1100–1200 m. Nei Mongol. margin serrate or crenate, apex rounded or sometimes acute. 20c. Viola collina var. glabricarpa K. Sun, Bull. Bot. Res., Flowers white, small; pedicels produced from leaf axils of stem, Harbin 14: 236. 1994. usually slightly exceeding or subequaling leaves, 2-bracteolate above middle; bracteoles linear. Sepals ovate or oblong, 3–4 × 光果球果堇菜 guang guo qiu guo jin cai 1.2–1.5 mm, apex acute, basal auricles orbicular, short, gla- Petioles and leaf blades puberulous. Capsules glabrous. brous or ciliate. Petals obovate, lateral ones 8–10 mm, glabrous or bearded, anterior one 7–8 mm, purple-violet striate, apex ● Forests, shaded and moist places; below 1400 m. Shandong emarginate; spur saccate, short, 1.2–1.7 mm, stout, ca. 2 mm in (Tai’an). diam., apex obtuse; anthers ca. 2.5 mm (appendage of con- 21. Viola hirta Linnaeus, Sp. Pl. 2: 934. 1753. nectives included); spur of 2 anterior stamens short, stout, apex obtuse and slightly curved. Ovary glabrous; styles slightly 硬毛堇菜 ying mao jin cai geniculate at base, slightly thickened upward; stigmas triangu- Herbs perennial, acaulescent, 5–15 cm tall. Rhizome lar, slightly thickened and narrowly margined on lateral sides brownish, robust, densely noded, often branched in upper part, and abaxially, apex conspicuously beaked, beak short, nearly sometimes with shortened, suberect flagellate branches. Leaves erect, stigma hole nearly as large as beak. Capsule ovoid. Fl. basal; stipules lanceolate, 1–2 cm, apex long acuminate, margin May–Jun, fr. Jun–Sep. 2n = 24.

86 VIOLACEAE

Mountain valleys, grasslands on mountain slopes, forest margins, Larix forests, river banks; 500–900 m. Heilongjiang, Jilin, Nei thickets, fields; 1900–4200 m. Gansu, Qinghai, Shaanxi, Sichuan, Xi- Mongol [Mongolia, Russia]. zang, Yunnan [Bhutan, N India, Nepal]. 25. Viola diamantiaca Nakai, Bot. Mag. (Tokyo) 33: 205. 23. Viola epipsiloides A. Löve & D. Löve, Bot. Not. 128: 516. 1919. 1976 [“1975”]. 大叶堇菜 da ye jin cai 溪堇菜 xi jin cai Viola diamantiaca f. glabrior (Kitagawa) Kitagawa; V. di- Viola repens Turczaninow ex Trautvetter & C. A. Meyer amantiaca var. glabrior Kitagawa. in Middendorff, Reise Sibir. 1(2,3) [Fl. Ochot.]: 18. 1856, not Schweinitz (1822). Herbs perennial, acaulescent, with long, slender stolons. Rhizome obliquely ascending or horizontal, slightly robust, Herbs perennial, acaulescent, 5–20 cm tall. Rhizome creep- densely noded, with numerous long, slender, brown roots. Basal ing, long and white, with remains of brown stipules. Leaves leaves 1, rarely 2 or 3, produced from top of rhizome; stipules basal, usually 2, sometimes 3; stipules free, white, ovate-lan- free, greenish, lanceolate or narrowly ovate-lanceolate, ca. 1 ceolate, ca. 8 mm, membranous, margin entire or sometimes 2- cm, submembranous when dry, margin remotely denticulate, or 3-denticulate, apex acuminate; petiole 2–5 cm at anthesis, to apex acuminate; petiole to 20 cm, slender, winged, usually 10 cm in fruit, very narrowly winged or wingless; leaf blade puberulous in upper part, glabrous in lower part; leaf blade broadly ovate, orbicular, or reniform, 1.5–2.5 × 2–3 cm, abaxially glaucous-green, adaxially green, cordate or ovate- abaxially ± puberulous, later glabrescent or sparsely puberulous cordate, 7–9 × 5–7 cm, thin, adaxially glabrous, puberulous only along veins, adaxially glabrous, base deeply cordate, mar- along veins, base shallowly or deeply cordate, margin obtusely gin shallowly crenate, apex acute or slightly obtuse. Flowers dentate, apex caudate-acuminate. Flowers purplish violet or purple or purplish; pedicels not or slightly exceeding leaves, glaucous, large, long pedicellate; pedicels solitary, slender, 2- robust, with 2 linear bracteoles in upper part. Sepals ovate-lan- bracteolate slightly above middle; bracteoles lanceolate, small. ceolate, ca. 4 mm, margin narrowly membranous, basal auri- Sepals ovate-lanceolate, glabrous, basal auricles short. Lateral cles short, apex truncate. Petals oblong-obovate, lateral ones petals 1.5–1.7 cm, glabrous, anterior one 1.8–2 cm (spur inclu- sparsely bearded, anterior one purple-striate, 1.5–1.8 cm (spur ded); spur short, ca. 4 mm, stout, apex obtuse. Capsule purple- included); spur short, ca. 3 mm, robust, horizontally straight or red punctate, ca. 1.3 cm. Fl. Apr–May, fr. Jun–Aug. slightly curved upward; spur of 2 anterior stamens shorter than anthers, ca. 3 mm. Ovary glabrous; styles slightly geniculate at Broad-leaved forests on mountains, forest margins, moist rocks; 600–1500 m. E Jilin, E Liaoning [Korea]. base, gradually thickened upward; stigmas thickly margined on lateral sides and at apex, very shortly beaked in front, with a 26. Viola diffusa Gingins in Candolle, Prodr. 1: 298. 1824. broad stigma hole at tip of beak. Capsule 5–10 mm, ellipsoid, 七星莲 glabrous, apex acuminate. Fl. May, fr. Jun. 2n = 24. qi xing lian Needle-leaved forests, in moist and boggy places, usually along Viola diffusa var. brevibarbata C. J. Wang; V. diffusa var. streams. 400–1300 m. Heilongjiang, Jilin, Nei Mongol, N Xinjiang brevisepala W. Becker; V. diffusa var. glabella H. Boissieu; V. [Japan, Korea, Russia; Europe, North America]. diffusa var. glaberrima W. Becker; V. diffusa subsp. tenuis In FRPS (51: 38. 1991), the name Viola epipsila Ledebour was (Bentham) W. Becker; V. diffusa var. tomentosa W. Becker; V. misapplied to this species. diffusoides C. J. Wang; V. tenuis Bentham; V. wilsonii W. Becker. 24. Viola brachyceras Turczaninow, Cat. Pl. Baical-Dahur. 191. 1839; Bull. Soc. Imp. Naturalistes Moscou 15: 301. 1842. Herbs annual, stiffly hairy or white puberulous throughout, or subglabrous, rarely glabrous, producing stolon at anthesis. 兴安圆叶堇菜 xing an yuan ye jin cai Stolon with rosulate leaves at top, usually producing adventi- Herbs perennial, acaulescent, 6–10 cm high. Rhizome 2–3 tious roots. Rhizome short, with numerous white rootlets and cm, oblique or vertical, rather sturdy, at top with remants of fibrous roots. Basal leaves numerous, fasciculate and rosulate, leaves and covered with rootlets. Leaves basal, few, 1–2 cm at or alternate on stolon; stipules adnate to petioles at base, 2/3 anthesis; petiole ca. 2 × as long as blade, slightly winged; leaf free, linear-lanceolate, 4–12 mm, margin remotely denticulate blade orbicular-cordate, both surfaces glabrous, margin ob- or fimbriate-dentate, apex acuminate; petiole 2–4.5 cm, con- scurely crenate, apex obtuse or rarely acute; leaves of fruiting spicuously winged, usually puberulous; leaf blade ovate or plant 3–5 × 3–5 cm, deeply cordate; stipules small, green but ovate-oblong, 1.5–3.5 × 1–2 cm, both surfaces densely white subsequently turning brown, ca. 1/2 adnate to petiole, margin puberulous on young leaves, later gradually sparsely so, but still sparsely fimbriate. Flowers white or light violet, ca. 8 mm; densely puberulous along veins and margin, or glabrous, base pedicel slightly exceeding leaves at anthesis. Sepals ovate-lan- broadly cuneate or truncate, rarely shallowly cordate, conspicu- ceolate, 3–4 mm, sparsely denticulate; sepal appendages short, ously decurrent to petiole, margin obtusely dentate and ciliate, apex rounded. Petals oblong-obovate; lateral petals bearded, apex obtuse or ± acute. Flowers purplish or yellowish, small, lower one shorter than others, lilac-striate; spur short, slightly long pedicellate, in basal leaf axils and leaf axils of stolon; exceeding sepal appendages. Style geniculate at base, flattened pedicels 1.5–8.5 cm, slender, glabrous or sparsely puberulous, on top. Capsule with dark brown dots, 6–10 mm. Fl. May–Jun, 2-bracteolate at middle; bracteoles linear. Sepals lanceolate, 4– fr. Jul–Sep. 2n = 20. 5.5 mm, apex acute, basal auricles short, margin sparsely cili-

VIOLACEAE 87

ate, apex rounded or remotely denticulate. Lateral petals obo- blade oblong-ovate or oblong, 1–2(–3) × 0.5–1.3 cm, both sur- vate or oblong-obovate, 6–8 mm, glabrous or shortly bearded, faces densely white villous, base cordate or rounded, margin anterior one ca. 6 mm (spur included), conspicuously shorter crenate, apex obtuse. Flowers light bluish violet; pedicels much than other petals, apex acute; spur very short, only ca. 1.5 mm, exceeding leaves, 3–4 cm, slender, sparsely puberulous, 2-brac- slightly exserted out of basal auricles of sepals; spur of 2 anteri- teolate above middle-bracteolate at middle; bracteoles oppo- or stamens triangular, short, broad. Ovary glabrous; styles cla- site, linear. Sepals narrowly lanceolate, 2.5–3 × ca. 1 mm, mar- vate, slightly geniculate, gradually thickened in upper part; stig- gin narrowly membranous, basal auricles short, only ca. 0.5 mas thickly margined on lateral sides and abaxially, slightly mm. Upper and lateral petals obovate, 1–1.1 cm, lateral ones raised in central part, shortly beaked in front. Capsule oblong, glabrous, anterior one boat-shaped, ca. 9 mm (spur included), 6–7 mm, ca. 3 mm in diam., glabrous, often with persistent styles. apex acute; spur ca. 1.5 mm. Ovary globose, glabrous; styles Seeds ca. 1 mm. Fl. Mar–May, fr. May–Oct. 2n = 26*, 74*. clavate, base geniculate, thickened at apex; stigmas narrowly margined on lateral sides, apex shortly beaked. Capsule ovoid- Mountain forests, forest margins, grassy slopes, stream valleys, orbicular, ca. 0.5 cm, glabrous. Fl. Feb–Apr and Aug–Sep. rock crevices; below 2000 m. Anhui, Chongqing, Fujian, SE Gansu, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangsu, ● Grassy places, forest margins, crevices; sea level to 1800 m. Jiangxi, S Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhu- Anhui, Fujian, Guangdong, Guizhou, Hubei, Hunan, Jiangxi. tan, Japan, India, Indonesia, Malaysia, Myanmar, Nepal, Papua New Guinea, Philippines, Thailand, Vietnam]. 29. Viola formosana Hayata, J. Coll. Sci. Imp. Univ. Tokyo 22: 28. 1906. 27. Viola grandisepala W. Becker, Bull. Misc. Inform. Kew 台湾堇菜 1928: 250. 1928. tai wan jin cai 阔萼堇菜 kuo e jin cai Herbs acaulescent. Rhizome erect, short, stout, usually with long stolons; stolons rooting at ends and producing dense Viola binchuanensis S. H. Huang; V. brunneostipulosa rosettes of leaves and flowers. Leaves basal; stipules adnate to Handel-Mazzetti. petioles only at base, free part lanceolate, margin fimbriate or Herbs perennial, dwarf, nearly acaulescent, 7–10 cm tall. erose; petioles slender, 1–15 cm, glabrous or sparsely puberu- Rhizome ca. 2 cm (shortened stem included), ca. 2 mm in lous; leaf blade abaxially greenish purple or glaucous, broadly diam., with dense nodes and leaves, sometimes with stolons. triangular-cordate or cordate to orbicular, 1–4 × 1–4 cm, both Leaves nearly basal; stipules adnate to petioles only at base, surfaces glabrous or sparsely puberulous, sometimes puberu- deep brown, upper ones ovate, inner ones broadly lanceolate, lous along teeth, base deeply cordate, margin crenate, apex ca. 1.2 cm, margin entire, apex acuminate; petiole 2–5 cm, soft, acute to rounded or obtuse. Flowers violet to white with dark- glabrous; leaf blade broadly ovate or suborbicular, 1–3 × 1.5–3 er veins, 1.5–2 cm in diam.; pedicels longer, sometimes to 15 cm, both surfaces glabrous, densely brown punctate, adaxially cm, glabrous or puberulous, 2-bracteolate above middle; bract- sparsely white puberulous near margin, base deeply cordate, eoles subulate. Sepals narrowly lanceolate, 3–5 × 0.5–1.5 mm, margin densely and shallowly crenulate, apex obtuse or orbicu- apex acuminate, basal auricles shorter, 0.5–1 mm, glabrous. lar. Flowers white, medium-sized; pedicels exceeding leaves, Upper petals subequaling lateral ones, ovate, ca. 1.2 cm, lat- glabrous, 2-bracteolate above middle; bracteoles brown, linear. eral ones glabrous, base cuneate, apex emarginate, anterior Sepals broadly ovate to ovate, ca. 5 × 3 mm, 3-veined, apex one larger, ca. 1.5 cm, apex deeply emarginate or shallowly 2- acute, basal auricles very short, abaxially brown punctate, mar- lobed; spur 5–7 mm, slightly curved. Styles nearly erect; stig- gin densely ciliate. Petals oblong-obovate, lateral ones glabrous, mas narrowly margined on lateral sides and abaxially, shortly anterior one ca. 1 cm (spur included); spur short, slightly longer beaked in front. Capsule globose to ellipsoid, 4–6 mm, gla- than basal auricles of sepals, 1.5–2 mm. Anthers of anterior brous. Seeds ovoid, ca. 1 mm. stamens ca. 2 mm, appendage of connectives ca. 1 mm; spur ● Mountain forests, mountains, grassy slopes; 1200–2500 m. Tai- short, only ca. 1.5 mm, broad. Ovary broadly ovoid; styles wan. nearly erect at base, slightly thickened in upper part; stigmas 1a. Leaf blade 1–3 × 1–3 cm, broadly curved forward and thickly beaked in front, with a large stigma cordate to orbicular, apex acute to hole at tip of beak. Fl. Apr and Sep, fr. May–Oct. rounded or obtuse, petiole 1–10 cm ... 29a. var. formosana ● Mountain slopes, shaded and moist places along roadsides; 1b. Leaf blade 1.5–4 × 1–4 cm, 1900–3000 m. Sichuan, Yunnan. triangular-cordate, apex acute, petiole to 15 cm ...... 29b. var. kawakamii 28. Viola lucens W. Becker, Bull. Misc. Inform. Kew 1928: 250. 1928. 29a. Viola formosana var. formosana 亮毛堇菜 liang mao jin cai 台湾堇菜(原变种) tai wan jin cai (yuan bian zhong) Herbs perennial, small, 5–7 cm tall, white villous through- Viola arisanensis W. Becker; V. formosana var. tozanensis out, acaulescent, stoloniferous. Rhizome erect, ca. 2 mm in (Hayata) C. F. Hsieh; V. taiwanensis W. Becker; V. tozanensis diam., densely noded, with numerous rootlets. Stolon slender, Hayata. often producing new plant at top. Leaves basal, rosulate; Petiole 1–10 cm; leaf blade 1–3 × 1–3 cm, broadly cor- stipules brown, lanceolate, margin fimbriate-dentate; petioles date to orbicular, apex acute to rounded or obtuse. Fl. Mar, fr. unequal in length, 0.2–2.5 cm, slender, densely villous; leaf May–Jun. 2n = 22*.

88 VIOLACEAE

● Mountains; 1400–2500 m. Taiwan. erect, brownish, slender. Stolon with clustered leaves at top. 29b. Viola formosana var. kawakamii (Hayata) Y. S. Chen Leaves basal, rosulate; stipules subulate, ca. 7 mm, margin & Q. E. Yang, comb. nov. erose, long fimbriate; petiole ca. 3 cm, slender, sparsely spread- ing puberulous; leaf blade ovate, elliptic, triangular-ovate to 川上氏堇菜 chuan shang shi jin cai lanceolate, 1–1.5 × 0.8–1 cm, abaxially glabrous or hairy, adax- ially with sparse white stiff hairs, base shallowly cordate, Basionym: Viola kawakamii Hayata, J. Coll. Sci. Imp. margin 8–10-crenate and ciliate, apex obtuse. Flowers pur- Univ. Tokyo 25(19): 52. 1908; V. formosana var. stenopetala plish, small; pedicels produced from axils of basal leaves and (Hayata) J. C. Wang et al.; V. hypoleuca Hayata; V. kawakamii those of leaves on stolon, much exceeding leaves, 8–10 cm, var. stenopetala Hayata; V. matsudae Hayata; V. takasagoensis slender, 2-bracteolate at middle; bracteoles subopposite, linear, Koidzumi. 5–7 mm, minutely ciliate. Sepals linear-lanceolate, 4–5 mm, Petiole to 15 cm; leaf blade triangular-cordate, 1.5–4 × 1– margin sparsely fimbriate-ciliate or glabrous, basal auricles 4 cm, apex acute. 2n = 22*. short. Petals obovate, upper ones subequaling lateral ones, 9–11 × ca. 4.5 mm, lateral petals glabrous or bearded, anterior petal ● Mountain forests, grassy slopes; 1200–2400 m. Taiwan. oblong, ca. 7.5 mm (spur included), apex acuminate; spur short, “Viola formosana var. kawakamii (Hayata) J. C. Wang” (Tax- ca. 1.5 mm; spur of 2 anterior stamens oblong, ca. 1.5 mm, onomic Study Viola Taiwan, 167. 1988) appeared in a Ph.D. disser- nearly as broad as anthers. Ovary ovoid, glabrous; styles nearly tation, which is not effectively published under Art. 30.5 of the Vienna erect, slender at base; stigmas slightly 2-lobed, lobes narrow Code. and slightly thick, slightly raised in central part, shortly beaked 30. Viola davidii Franchet, Nouv. Arch. Mus. Hist. Nat., sér. 2, in front. Capsule ca. 5 mm. 8: 203. 1886. ● Grassy slopes; below 1100 m. Taiwan. 深圆齿堇菜 shen yuan chi jin cai 1a. Leaf blade ovate or elliptic, apex obtuse, abaxially glabrous; sepals sparsely Viola davidii var. paucicrenata W. Becker; V. schneideri fimbriate-ciliate along margin; lateral W. Becker; V. smithiana W. Becker. petals beardless ...... 31a. var. nagasawae Herbs perennial, nearly acaulescent, 7–10 cm tall, glabrous. 1b. Leaf blades triangular-ovate to lanceolate, Rhizome obliquely ascending, shortly noded, with dense root- apex acute, both surfaces stiffly hairy; lets. Stolon developed, 10–15 cm, with scattered leaves and flow- sepals sparsely ciliate or glabrous ers, producing adventitious roots at nodes, top usually develop- along margin; lateral petals bearded ...... 31b. var. pricei ing into a new plant. Leaves nearly basal; stipules mostly free, brown, broadly lanceolate, 1–1.5 cm, adaxially brown striate, 31a. Viola nagasawae var. nagasawae margin remotely fimbriate-dentate, apex long acuminate; peti- 台北堇菜(原变种) tai bei jin cai (yuan bian zhong) oles unequal in length, longest to 5 cm; leaf blade abaxially often glaucous, adaxially greenish, ovate or ovate-orbicular, 2– Viola acutilabella Hayata; V. nagasawae var. acutilabella 7 × 1.5–3.5 cm, both surfaces glabrous, with brown glandular (Hayata) Nakai. dots when dry, base deeply cordate, margin shallowly 6–8- Leaf blade ovate or elliptic, apex obtuse, abaxially gla- crenate on each side, apex rounded or acute. Flowers white or brous; sepals sparsely fimbriate-ciliate along margin; lateral purplish; pedicels exceeding or subequaling leaves, 2-bracteo- petals beardless. Fl. Mar–Nov, fr. May–Nov. 2n = 48*. late above middle; bracteoles linear. Sepals lanceolate or ovate- lanceolate, 5–6 × 1.5–2 mm, apex acute, basal auricles short, ● Moist places on grassy slopes; 200–1100 m. Taiwan. margin narrowly membranous, apex truncate. Petals oblong-obo- 31b. Viola nagasawae var. pricei (W. Becker) J. C. Wang & vate, 7–8 mm, lateral ones bearded, anterior one shorter; spur T. C. Huang, Taiwania 35: 47. 1990. saccate, short, 1.5–2 mm, ca. 2 mm in diam.; spur of anterior stamens oblong, short, subequaling anthers, ca. 2 mm. Ovary 锐叶台北堇菜 rui ye tai bei jin cai oblong, glabrous; styles clavate, base nearly erect, gradually Viola pricei W. Becker, Bull. Misc. Inform. Kew 1928: thickened upward; stigmas conspicuously and broadly mar- 252. 1928. gined on lateral sides, beaked in front, beak upward and straight, short, stout, with large stigma hole at tip. Capsule oblong, 5–7 Leaf blade triangular-ovate to lanceolate, both surfaces mm, glabrous. Fl. Mar–May and Sep, fr. May–Oct. stiffly hairy, apex acute. Sepals sparsely ciliate or glabrous along margin. Lateral petals bearded. 2n = 48*. ● Mountain forests, forest margins, grassy slopes, stream valleys, roadsides; 1200–2800 m. Chongqing, Fujian, Guangdong, Guangxi, Gui- ● Grassy slopes; below 500 m. Taiwan. zhou, Hubei, Hunan, Jiangxi, Sichuan, SE Xizang, Yunnan, Zhejiang. 32. Viola sikkimensis W. Becker, Beih. Bot. Centralbl., Abt. 2, 31. Viola nagasawae Makino & Hayata, J. Coll. Sci. Imp. Univ. 34: 260. 1916. Tokyo 22: 30. 1906 [“nagasawai”]. 锡金堇菜 xi jin jin cai 台北堇菜 tai bei jin cai Viola hookeri Thomson & W. Becker (1916), not Franchet Herbs perennial, acaulescent, with slender stolon. Rhizome (1886).

VIOLACEAE 89

Herbs perennial, acaulescent. Rhizome short, 2–2.5 cm, Herbs perennial, acaulescent, with stolons. Rhizome short, stout, 4–7 mm in diam., densely noded, often with brown re- 2–4 cm, stout, 2–3 mm in diam., densely noded, often with mains of stipules. Stolon to more than 10 cm; internodes longer, brown remains of stipules. Stolon to 20 cm, internodes longer, producing rootlets at nodes. Leaves basal; stipules brown, lan- producing rootlets at nodes. Leaves basal; stipules brown, lan- ceolate, 1–1.3 cm, margin long fimbriate-dentate; petiole 3–10 ceolate, 8–10 mm, margin long or sparsely fimbriate; petiole cm, glabrous; leaf blade abaxially gray-green, adaxially deep 1.5–12 cm, glabrous; leaf blade abaxially gray-green, adaxially green and nitid, broadly ovate or suborbicular, 1.5–5 × 1.5–4 deep green, broadly ovate or suborbicular, 1.5–4 × 1.3–3.5 cm, cm, both surfaces glabrous or sparsely puberulous, base deeply dotlike brown glandular, abaxially glabrous, adaxially sparsely cordate, margin densely and shallowly crenate, apex obtuse. puberulous and scabrous, base deeply cordate, margin densely Flowers white or purplish; pedicels exceeding leaves, glabrous, and shallowly rounded-serrate, apex obtuse; leaves in stolons 2-bracteolate above middle; bracteoles subopposite, linear, mar- smaller. Flowers white; pedicels 2–2.5 cm, glabrous, 2-bracteo- gin remotely denticulate. Sepals narrowly lanceolate, 5–6 mm, late at middle; bracteoles subopposite, linear, margin entire. Se- glabrous, 3-veined, basal auricles rounded or remotely dentate pals linear-lanceolate, ca. 3 × 1.2 mm, puberulous or nearly gla- at apex. Petals oblong-obovate, 1–1.2 cm × 4–4.5 mm, lateral brous when mature, basal auricles ca. 1 mm, apex acute. Upper ones glabrous, anterior one purple veined, shorter, 1–1.2 cm petals spatulate, ca. 7 × 4 mm, lateral ones spatulate, ca. 8 × 3 (spur included); spur ca. 2 mm, ca. 2.5 mm in diam., slightly mm, glabrous, anterior one oblong, ca. 9 × 3 mm; spur ca. 1 enlarged; spur of 2 anterior stamens angular, slightly longer mm, ca. 5 mm in diam. Capsules ovoid, 5–6 mm. Seeds glo- than anthers, apex obtuse. Ovary conic, glabrous; styles clavate, bose, ca. 1.2 mm in diam. Fl. Mar–Apr, fr. Aug–Sep. gradually thickened upward; stigmas flat at apices, broadly mar- gined on lateral sides and abaxially, shortly beaked at apex, ● Forests; 1600–2700 m. Yunnan. beak straight. Capsule ovoid-orbicular. Fl. Apr–May, fr. May– 35. Viola yunnanensis W. Becker & H. Boissieu, Bull. Herb. Jun. Boissier, sér. 2, 8: 740. 1908. Mountain forests, forest margins, streamsides, riversides; 1500– 云南堇菜 yun nan jin cai 2500 m. SE Xizang, Yunnan [?Bhutan, India, Myanmar, Nepal]. Herbs perennial. Rhizome elongate, obliquely ascending 33. Viola kwangtungensis Melchior, Sunyatsenia 1: 124. 1933. or procumbent and stemlike, with adventitious roots at nodes. 广东堇菜 guang dong jin cai Stem absent or shorter, less than 2 cm. Stolon to 37 cm, slender, usually densely white puberulous, with clustered leaves at top Viola sikkimensis W. Becker var. debilis W. Becker. and often producing a new plant. Leaves nearly basal or alter- Herbs perennial, small, acaulescent or nearly so, 4–9 cm nate on stolon; stipules mostly free, glaucous, lanceolate, 1–1.3 tall, glabrous, sometimes stoloniferous. Rhizome nearly erect, cm, membranous, margin fimbriate-dentate, apex narrowly long slender, densely noded. Leaves basal; stipules free or adnate acuminate; petioles unequal in length, 3–8 cm, densely spread- to petioles only at base, brown, lanceolate, ca. 0.5 mm, mar- ing white puberulous; leaf blade abaxially gray-green, adaxially gin fimbriate, apex acuminate; petioles unequal in length, 2–5 deep or dark green, oblong or oblong-ovate, 3–8 × 2–4 cm, cm, glabrous; leaf blade abaxially purple, adaxially deep green, broadest near middle, both surfaces densely gray-white puberu- orbicular or sometimes reniform, 1–3 × 1–3 cm, both surfaces lous, more densely so on young leaves, base shallowly and nar- glabrous, base shallowly cordate or truncate, margin deeply cre- rowly cordate, margin large crenate, apex acute or acuminate. nate, shortly spinose in between teeth, apex obtuse. Flowers Flowers white, 1.5–1.7 cm, ca. 1.5 cm in diam.; pedicels of light violet, sometimes white; pedicels 4–9 mm, slender, 2- cleistogamous flowers produced from basal leaf axils much bracteolate at upper part; bracteoles linear. Sepals lanceolate, shorter than leaves, those on stolon much longer than leaves; usually pubescent, 3–5 × 1.5–2 mm, apex ± acute, basal auri- pedicels densely puberulous, 2-bracteolate above middle; bract- cles absent, margin membranous, apex truncate. Petals obovate- eoles opposite, linear, 8–10 mm. Sepals linear-lanceolate or lan- oblong, upper ones 1–1.2 cm × ca. 4 mm, lateral ones subequal- ceolate, 5–7 mm, basal auricles short, sparsely white puberu- ing upper ones, glabrous, anterior one shorter, ca. 9 mm (spur lous along veins, 3-veined, margin densely ciliate. Upper pe- included), purple veined; spur saccate, shorter, ca. 2 mm. An- tals oblong, 1.3–1.5 cm × 5–6.5 mm, lateral ones ca. 1 cm, thers ca. 1.5 mm. Ovary globose, with brown glandular dots; usually shortly bearded, anterior one shorter, deeply veined, styles clavate, base geniculate; stigmas narrowly margined on 8–9 mm (spur included), apex acute; spur saccate, very short, lateral sides and abaxially, shortly beaked in front. Capsule ca. 1.5 mm, slightly curved downward; spur of 2 anterior sta- ellipsoid, ca. 7 mm, glabrous, often with brown glandular dots. mens very shortly angular. Ovary ovoid-globose, glabrous; Fl. Apr–May, fr. May–Aug. styles clavate, base nearly erect, gradually thickened upward; stigmas narrowly margined, shortly beaked in front, beak ● Forests, forest margins, grasslands on mountain slopes, stream straight, with a smaller stigma hole at tip. Capsule green, ob- valleys, shaded places on rocks; 600–2000 m. Fujian, N Guangdong, Hunan, Jiangxi, Sichuan. long or subglobose, small, 5–7 mm. Seeds globose. Fl. Feb– Jun, fr. May–Dec. 34. Viola duclouxii W. Becker, Bull. Misc. Inform. Kew 1928: 249. 1928. Mountain forests, grasslands along forest margins, moist places on rocks along streamsides and roadsides; 1300–2400 m. Hainan, S Yun- 紫点堇菜 zi dian jin cai nan [Indonesia, Malaysia, Myanmar, Vietnam].

90 VIOLACEAE

36. Viola fargesii H. Boissieu, Bull. Herb. Boissier, sér. 2, 2: slightly shorter than leaves, 2-bracteolate above middle; 333. 1902. bracteoles linear. Sepals lanceolate, ca. 5 × 1–1.2 mm, glabrous, margin narrowly membranous, basal auricles short, less than 1 柔毛堇菜 rou mao jin cai mm, apex truncate, sparsely shallowly dentate. Petals obovate, Viola adenothrix Hayata; V. adenothrix var. tsugitakaensis upper ones subequaling lateral ones, ca. 1.3 × 0.5 cm, lateral (Masamune) J. C. Wang & T. C. Huang; V. arisanensis W. ones glabrous, anterior one shorter, ca. 1.1 cm × ca. 3 mm (spur Becker; V. brachycentra Hayata; V. canescens Wallich f. gla- included); spur saccate, short, 2–2.5 mm, stout, 2–2.5 mm in brescens W. Becker; V. canescens subsp. lanuginosa W. Beck- diam.; spur of 2 anterior stamens acutangular, ca. 2 mm, sub- er; V. principis H. Boissieu; V. principis var. acutifolia C. J. equaling anthers. Ovary oblong-globose, glabrous; styles cla- Wang; V. pulla W. Becker; V. thrichopoda Hayata; V. tsugitaka- vate, base slightly geniculate, gradually thickened upward; ensis Masamune. stigmas slightly raised at apex, thickened and narrowly mar- gined on lateral sides, inconspicuously beaked at apex, with a Herbs perennial, spreading white puberulous throughout. small stigma hole open upward at tip of beak. Capsule ellipsoid, Rhizome robust, 2–4 cm, 3–7 mm in diam. Stolon longer, ca. 6 mm, glabrous. Fl. Mar–Apr, fr. May–Jun. elongated, puberulous, sometimes stemlike. Leaves nearly basal or alternate on stolon; stipules mostly free, brown or greenish, ● Mountain forests, forest margins, grasslands; 1300–1900 m. dark striate, broadly lanceolate, 1.2–1.8 cm × 3–4 mm, margin Guangxi, Guizhou, Sichuan, Yunnan. long fimbriate-dentate, apex acuminate; petiole 5–13 cm, 38. Viola thomsonii Oudemans in Miquel, Ann. Mus. Bot. densely villous, wingless; leaf blade ovate or broadly ovate, Lugduno-Batavi 3: 74. 1867. sometimes suborbicular, 2–6 × 2–4.5 cm, abaxially puberulous, 毛堇菜 more densely so along veins, base broadly cordate or narrowly mao jin cai cordate, margin densely and shallowly obtusely dentate, apex Herbs perennial, acaulescent. Rhizome long, longest to 10 rounded, rarely acuminate. Flowers white; pedicels usually ex- cm, stout, 4–6 mm in diam.; internodes 2–4 cm. Stolon long, to ceeding leaves, densely spreading white puberulous, 2-bracteo- 15 cm, slender, flagellate, with alternate small leaves, often late above middle; bracteoles opposite, linear. Sepals narrowly producing new plant at top. Leaves basal; stipules deep brown, ovate-lanceolate or lanceolate, 7–9 mm, apex acuminate, basal lanceolate or broadly lanceolate, 1–1.5 cm, dry membranous, auricles short, ca. 2 mm, 3-veined, margin puberulous outside, margin fimbriate-dentate, glabrous; petioles unequal in length, apex obtuse. Petals oblong-obovate, 1–1.5 cm, apex ± acute, 6–11 cm, very narrowly winged in upper part, glabrous; lower lateral ones slightly bearded, anterior one shorter, ca. 7 mm leaves cordate, smaller, upper ones larger, ovate-cordate, 6–8 × (spur included), apex acute; spur saccate, short, 2–2.5 mm, 4–4.5 cm, both surfaces sparsely white puberulous or glabrous stout, ca. 2 mm in diam.; spur of 2 anterior stamens angular, adaxially, base deeply cordate, margin densely and shallowly slightly longer than anthers, apex acute. Ovary conic, glabrous; crenate, apex acuminate. Flowers white or light violet; pedicels styles clavate, base slightly geniculate, thickened upward, slight- usually not exceeding leaves, glabrous, 2-bracteolate tightly ly flat at apex, conspicuously margined on lateral sides, shortly near flower; bracteoles brown, linear, 1–1.3 cm, dry mem- beaked in front, with a stigma hole open upward at tip of beak. branous. Sepals lanceolate, 4.5–5.5 × 1.2–1.8 mm, apex acute, Capsule narrowly orbicular, ca. 8 mm, glabrous. Fl. Mar–Jun, basal auricles longer, ca. 2 mm, apex shallowly 2- or 3-dentate, fr. Jun–Sep. 2n = 24*. sparsely ciliate, 3-veined. Petals oblong-obovate, ca. 1.5 cm, ● Mountain forests, forest margins, grasslands, stream valleys, lateral ones shortly bearded, anterior one shorter, inside purple roadsides; 600–3800 m. S Anhui, Fujian, N Guangdong, N Guangxi, striate, shortly spurred; spur saccate, shorter than basal auricles Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, of sepals, ca. 2 mm in diam.; spur of 2 anterior stamens shortly Zhejiang. angular, subequaling anthers. Ovary ovoid-globose, glabrous; styles clavate; stigmas narrowly margined on lateral sides, 37. Viola mucronulifera Handel-Mazzetti, Sinensia 2: 4. 1931. shortly beaked in front, beak erect, with smaller stigma hole at 小尖堇菜 xiao jian jin cai tip. Capsule ellipsoid. Seeds dark brown, globose, 7–10 mm. Fl. Apr–Jun, fr. Jul–Aug. Herbs perennial, nearly acaulescent, or stem short, only ca. 1 cm. Rhizome long, slender; internodes remote. Stolon to 15 Mountain forests, forest margins, stream valleys, shaded and cm, slender, glabrous, producing numerous adventitious roots at moist places along roadsides; 800–2400 m. SE Xizang, NW and S Yun- nodes, top developing into a new plant. Leaves nearly basal or nan [Bhutan, E India, Myanmar, Nepal]. clustered on shortened stem; stipules adnate to petioles only at 39. Viola sumatrana Miquel, Fl. Ned. Ind., Eerste Bijv. 389. base, free part lanceolate, brown striate, margin long fimbriate- 1861. dentate; petioles subequaling blades, to 7 cm, glabrous; leaf blade abaxially gray-green, adaxially deep green, ovate-cordate 光叶堇菜 guang ye jin cai or elliptic-cordate, 2–5 × 1.5–3 cm, both surfaces glabrous, Viola hossei W. Becker. abaxially often sparsely spinose along raised midvein, base nar- rowly cordate, margin serrate, 6–10-toothed on each side, teeth Herbs perennial, acaulescent. Rhizome 1–20 cm, 1.5–2.5 suborbicular, conspicuously spinose at apices, spines spreading mm in diam., with brown remains of stipules at nodes. Stolon or bending forward and often reddish glandular at apices, apex 15–20 cm, longest to 40 cm, slender, glabrous, producing ad- acute. Flowers white or purplish; pedicels subequaling or ventitious roots. Leaves basal or alternate on stolon; stipules

VIOLACEAE 91

free, deep brown, linear-lanceolate, 7–15 mm, margin long rootlets at nodes. Leaves basal; stipules free, greenish, lanceo- fimbriate-dentate; petioles unequal in length, 2–9 cm, narrowly late or narrowly ovate, 5–8 mm, submembranous when dry, winged in upper part, puberulous or glabrescent; leaf blade margin entire or remotely denticulate, apex acuminate; petiole abaxially greenish, adaxially deep green or dark green, triangu- soft, very narrowly winged, to 14 cm after anthesis; leaf blade lar-ovate or oblong-ovate, 2–5 × 1.5–3 cm, usually broadest abaxially greenish, adaxially green, broadly ovate or subreni- near base, both surfaces glabrous or sparsely white puberulous form, 2–6 × 2–5 cm, abaxially densely white puberulous, grad- and dotlike brown glandular, base deeply cordate, margin shal- ually less hairy on old leaves, adaxially sparsely white puberu- lowly serrate or shallowly crenate, glandular at apices of teeth, lous at base and along margin, base shallowly cordate, rarely apex long acuminate. Flowers violet to light purple or white; deeply cordate, auricles often involute, margin densely serru- pedicels usually not exceeding leaves, 2-bracteolate at or slight- late, apex acuminate. Flowers purple, large, long pedicellate; ly above middle; bracteoles opposite, rarely alternate, 3–8 mm. pedicels subequaling leaves, glabrous, 2-bracteolate slightly Sepals linear-lanceolate, 5–6 mm, apex acute; appendages very above middle; bracteoles opposite, lanceolate. Sepals ovate or short, only ca. 0.5 mm, 3-veined, dotlike rusty glandular, mar- oblong-ovate, ca. 7 mm, glabrous, basal auricles short, apex ob- gin membranous, apex truncate. Petals oblong-ovate, 8–10 × ca. tuse or truncate, remotely dentate. Petals obovate, lateral ones 3 mm, lateral ones glabrous, anterior one shorter, ca. 7 mm sparsely bearded or glabrous, anterior one spatulate, 1.8–2 cm (spur included); spur saccate, short, ca. 1.5 mm, usually twice (spur included); spur saccate, 3–4 mm, ca. 4 mm in diam.; spur as long as calycine appendages; spur of 2 anterior stamens of 2 anterior stamens shorter, subequaling anthers, ca. 3 mm, short, wide, shortly angular, usually subequaling anthers. Ovary ca. 1.5 mm in diam., apex slender. Ovary glabrous; styles ovoid-globose; styles clavate, base slightly geniculate, gradu- slightly geniculate at base, slightly thickly margined on lateral ally thickened upward; stigmas thickened and conspicuously sides, slightly flat in central part, shortly beaked in front. Cap- margined on lateral sides, margin spreading, shortly beaked in sule ellipsoid, large, ca. 1.2 cm, glabrous, apex mucronate. front, beak erect, with a smaller stigma hole at tip. Capsule Seeds glaucous, ovoid-globose, ca. 2.8 mm, ca. 1.8 mm in ellipsoid, small, 5–7 mm, dotlike brown rusty glandular. Seeds diam., smooth. Fl. Apr–May, fr. Jun–Aug. 2n = 24. globose, small, ca. 0.5 mm in diam. Fl. spring–summer, fr. summer–autumn. 2n = 46. Mixed needle-leaved and broad-leaved forests, broad-leaved for- ests, forest margins, thickets, grasslands on mountain slopes; 100–1300 Shaded forests, forest margins, streamsides, rock crevices along m. Anhui, Hunan, Jiangxi, Liaoning, Shandong, Zhejiang [Japan, Korea]. streamsides; below 2400 m. Guangxi, Guizhou, Hainan, S Yunnan [In- donesia, Malaysia, Myanmar, Thailand, Vietnam]. 42. Viola moupinensis Franchet, Bull. Soc. Bot. France 33: 412. 1886. 40. Viola nuda W. Becker, Repert. Spec. Nov. Regni Veg. 26: 26. 1929. 萱 xuan 裸堇菜 luo jin cai Viola mairei H. Léveillé; V. moupinensis var. lijiangensis C. J. Wang; V. palustris var. moupinensis (Franchet) Franchet; Herbs perennial, acaulescent, stoloniferous, or with very V. paravaginata H. Hara; V. rosthornii E. Pritzel; V. vaginata short stem, 7–13 cm tall. Rhizome erect, 1–3 cm, ca. 3 mm in Maximowicz subsp. alata W. Becker; V. vaginata var. sutchu- diam., conspicuously noded; stolons 1 or several, 10–15 cm, ensis Franchet ex H. Boissieu. with leaves at top. Stipules free, lanceolate, 1–2 cm, margin re- motely fimbriate-dentate, apex acuminate; petiole 2.5–8 cm, Herbs perennial, acaulescent, sometimes with ascending wingless; leaf blade ovate, 2–5 × 2.5–4 cm, but smaller on sto- stolon to 30 cm, clustered with several leaves at top of stolon. lons, both surfaces glabrous, base broadly cordate, margin shal- Rhizome erect or sometimes obliquely ascending, to 15 cm, 6– lowly serrate, apex ± acute. Flowers white, purplish striate and 10 mm in diam., densely noded, usually with brown remains of punctate at base; pedicels 2-bracteolate at upper part; bracteoles stipules and dense rootlets. Leaves basal; stipules free, brown- 6–8 mm. Sepals narrowly ovate, ca. 5 × 2–2.5 mm, basal auri- ish or light colored in upper half, ovate, 1–1.8 cm, margin re- cles short, margin denticulate. Upper petals obovate, lateral ones motely denticulate or entire, apex acuminate; petiole winged, 4– obovate-oblong, ca. 1 cm, inside shortly bearded, anterior one 10 cm, to 25 cm after anthesis; leaf blade cordate or reniform- shorter, ca. 8 mm, apex ± acute; spur of 2 anterior stamens cordate, 2.5–5 × 3–4.5 cm, accrescent after anthesis and reni- acutely angular, ca. 1.5 mm. Anthers ca. 2 mm, appendage of form, ca. 9 × 10 cm, both surfaces glabrous, sometimes slightly connectives ca. 1.5 mm. Styles geniculate at base; stigmas flat puberulous only along veins abaxially, base narrowly sinuate or at top, narrowly margined on lateral sides and abaxially, very broadly triangular, lateral auricles often involute at anthesis, shortly beaked in front. Capsules unknown. Fl. Apr. margin glandular obtusely dentate, apex acute or acuminate. Flowers purplish or white, purple-striate, large; pedicels not ● Mountain thickets, streamsides, riversides; ca. 2700 m. W Yunnan. exceeding leaves, 2-bracteolate at middle; bracteoles linear. Sepals lanceolate or narrowly ovate, apex ± acute, basal auri- 41. Viola rossii Hemsley, J. Linn. Soc., Bot. 23: 54. 1886. cles short, apex truncate, remotely and shallowly dentate, mar- 辽宁堇菜 liao ning jin cai gin narrowly membranous. Petals oblong, lateral ones bearded, anterior one ca. 1.5 cm (spur included); spur saccate, conspicu- Herbs perennial, acaulescent. Rhizome erect or obliquely ously longer than basal auricles of sepals, stout; spur of 2 an- ascending, deep brown, 2–4 cm, 3–4 mm in diam., sometimes terior stamens ca. 1 mm, ca. 1.1 mm in diam., apex obtuse. branched in upper part, densely noded, with numerous brown Ovary glabrous; styles slightly geniculate forward at base,

92 VIOLACEAE

thickened in upper part; stigmas truncate, thickly margined on m. N Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol [Mongolia, Rus- lateral sides and abaxially, shortly beaked in front, beak hori- sia]. zontal. Capsule ellipsoid, ca. 1.5 cm, glabrous, brown glandu- 44. Viola dissecta Ledebour, Fl. Altaic. 1: 255. 1829. lar. Seeds large, obovoid, ca. 2.5 mm, ca. 2 mm in diam., base acute, apex rounded. Fl. Mar–May, fr. May–Jul. 裂叶堇菜 lie ye jin cai

Open places at forest margins, thickets, streamsides, grassy slopes; Herbs perennial, acaulescent, highly variable in height, 3– 600–3600 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, 17 cm tall at anthesis, 4–34 cm tall at fruiting. Rhizome erect, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Sichuan, Xizang, Yunnan, short, 5–12 mm, 3–8 mm in diam., densely noded, often pro- Zhejiang [Bhutan, India (Sikkim), Nepal]. ducing from lower part several thicker yellowish roots. Stipules Wang (Bull. Bot. Res., Harbin 8(2): 20. 1988) described a yellow- of basal leaves glaucous to greenish, over 2/3 adnate to petioles, flowered variety, Viola moupinensis var. lijiangensis, and stated that it free part narrowly lanceolate, submembranous, margin remote- differs from the typical variety in having yellow flowers. According to ly denticulate, apex acuminate; petioles highly variable in length our examination of the type material and field work at the type locality, and indumentum, 1.5–24 cm, those of young leaves often pu- we consider that the flower color of the type specimens might have been bescent, glabrescent; blades of basal leaves orbicular, reniform, wrongly recorded. or broadly ovate, 1.2–9 × 1.5–10 cm, both surfaces white pu- 43. Viola dactyloides Roemer & Schultes, Syst. Veg. 5: 351. bescent, later glabrescent or sparsely pubescent only adaxially, 1819. veins conspicuously raised and pubescent or glabrous abaxially, usually 3-, rarely 5-sect, segments linear, oblong, or narrowly 掌叶堇菜 zhang ye jin cai ovate-lanceolate, 0.2–3 cm wide, margin entire or sparsely ir- Viola dactyloides var. multipartita W. Becker. regularly dentate, or subpinnatilobed, terminal lobes usually mi- nutely ciliate, blade not divided, margin incised. Flowers pur- Herbs perennial, acaulescent, 7–20 cm tall. Rhizome plish or purple-violet, large; pedicels usually subequaling or slightly obliquely ascending, short, 6–20 mm, with numerous slightly exceeding leaves, usually shorter than leaves at fruiting, red-brown roots. Leaves basal, long petiolate; stipules greenish, hairy or glabrous, 2-bracteolate above middle; bracteoles linear. ovate-lanceolate, over 1/2 adnate to petioles, free part long Sepals ovate, oblong-ovate, or lanceolate, 4–7 mm, 3-veined, acuminate at apex, margin entire or remotely fimbriate-denticu- margin narrowly membranous, apex ± acute, basal auricles short, late, dry membranous; petiole to 15 cm, usually white puberu- 1–1.5 mm, apex truncate, entire or 1- or 2-denticulate. Upper lous in lower part, glabrescent; leaf blade 5-palmatisect, seg- petals narrowly obovate, 8–13 × 6–9 mm, slightly curved up- ments oblong, oblong-ovate, or broadly lanceolate, 3–4 × 0.5–1 ward in upper part, lateral ones oblong-obovate, 7–10 mm × ca. cm (slightly accrescent at fruiting), abaxially more densely pu- 6 mm, sparsely bearded, anterior one (including spur) 1.4– 2.2 berulous along veins and margin, adaxially glabrous or sparse- cm; spur conspicuous, cylindric, 4–8 mm, 2–3 mm in diam., ly puberulous, base acuminate and shortly stipitate, stipes white apex obtuse and slightly enlarged. Anthers 1.5–2 mm, append- puberulous or glabrous, margin remotely serrate or slightly re- age of connectives 1.5–2 mm, spur of anterior 2 stamens long, pand, or shallowly emarginate-dentate, sometimes some seg- 3–5 mm, slender, 0.5–0.7 mm in diam. Ovary ovoid-globose, ments shallowly or deeply 2- or 3-lobed, apex ± acute. Flowers ca. 1.8 mm, glabrous; styles clavate, 2–2.5 mm, slightly slender purplish, large, fragrant, long pedicellate; pedicels deep green, at base and slightly geniculate forward; stigmas slightly thick- usually not exceeding leaves, glabrous, 2-bracteolate below ened and margined on lateral sides and abaxially, margins spread- middle; bracteoles linear, small, 5–8 mm, margin entire or few ing, shortly beaked in front, with a conspicuous stigma hole at denticulate. Sepals oblong or lanceolate, ca. 8 mm, apex slight- tip of beak. Capsule oblong or ellipsoid, 7–18 mm, apex acute, ly obtuse, basal auricles short, only 1–1.5 mm, 3-veined, margin pericarp hard, glabrous. Fl. Apr–Sep, fr. May–Oct. narrowly membranous, apex truncate. Upper petals broadly obo- vate, ca. 1.6 × 0.85 cm, claw ca. 4.5 mm; lateral petals oblong- Grasslands in logged forest areas and on mountain slopes, mixed forest margins, thickets, field margins, roadsides; below 3000 m. Gansu, obovate, ca. 1.5 × 0.7 cm, conspicuously bearded; anterior petal Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, obovate, 2–2.3 cm (spur included), spur long, 5–6 mm, stout, Shaanxi, Shandong, Shanxi, N Sichuan [Korea, Mongolia, Russia]. ca. 2.5 mm in diam., slightly curved upward, apex obtuse. An- thers ca. 2 mm, appendage of connectives ca. 2 mm, spur of 2 1a. Leaf blade deeply divided ...... 44a. var. dissecta anterior stamens long, ca. 4 mm, slender, 0.2–0.6 mm in diam. 1b. Leaf blade margin incised ...... 44b. var. incisa Ovary ovoid-globose, ca. 2 mm, glabrous; styles slender at base 44a. Viola dissecta var. dissecta and geniculate forward, conspicuously thickened in upper part; stigmas 2-lobed, slightly thickened and narrowly margined on 裂叶堇菜(原变种) lie ye jin cai (yuan bian zhong) lateral sides, margin erect, slightly concave in central part, short- ly beaked in front, beak obliquely ascending and larger, with a Viola dissecta f. pubescens (Regel) Kitagawa; V. dissecta small stigma hole at tip. Capsule purplish (immature), ellipsoid, var. pubescens (Regel) Kitagawa; V. ×lii Kitagawa; V. pinnata ca. 7 mm, glabrous, apex acute. Seeds brown-red, ovoid-globose, Linnaeusvar.dissecta(Ledebour) Regel;V.pinnata subsp. multi- ca. 2.5 mm, ca. 2 mm in diam. Fl. and fr. May–Aug. 2n = 24. fida W. Becker.

Deciduous broad-leaved forests, mixed needle-leaved and broad- Leaf blade cordate, glabrous or sparsely pubescent, deeply leaved forests, forest margins, thickets, shaded rock crevices; 500–700 divided. Fl. Apr–Sep, fr. May–Oct. 2n = 24*.

VIOLACEAE 93

Grasslands on mountain slopes, mixed forest margins, thickets, stigma hole at tip of beak. Capsule narrowly ellipsoid, large, 1– field margins, roadsides; below 3000 m. Gansu, Hebei, Heilongjiang, 1.6 cm, glabrous, apex acute. Seeds numerous, ovoid, ca. 2.2 Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong, mm, ca. 1.5 mm in diam. Shanxi, N Sichuan [Korea, Mongolia, Russia]. Mixed forests, forest margins, shaded and moist places in stream 44b. Viola dissecta var. incisa (Turczaninow) Y. S. Chen, comb. valleys, thickets on sunny slopes, grassy slopes; below 2000 m. Anhui, et stat. nov. Chongqing, Hebei, Henan, Hubei, Jiangsu, Jiangxi, Liaoning, Shan- dong, Zhejiang [Japan, Korea, SE Russia]. 总裂叶堇菜 zong lie ye jin cai 1a. Terminal lobe of leaves irregularly Basionym: Viola incisa Turczaninow, Bull. Soc. Imp. dentate or shallowly divided .... 45a. var. chaerophylloides Naturalistes Moscou 15: 302. 1842; V. fissifolia Kitagawa; V. 1b. Terminal lobe of leaves triparted ×jettmarii Handel-Mazzetti. to pinnatiparted or 3-lobed ...... 45b. var. sieboldiana Leaf blade ovate, 1.5–3 × 1–1.5 cm, accrescent at fruiting, 45a. Viola chaerophylloides var. chaerophylloides to 5 cm, both surfaces densely white puberulous, base broadly cuneate, margin irregularly emarginate-lobed to emarginate-fid, 南山堇菜(原变种) nan shan jin cai (yuan bian zhong) lower lobes irregularly 2- or 3-dentate, apex ± acute. Fl. Apr– May, fr. Jun–Oct. Viola pinnata Linnaeus var. chaerophylloides Regel, Pl. Radd. 1: 222. 1861; V. albida Palibin var. chaerophylloides Forest margins on mountains, grasslands in logged forest areas, (Regel) F. Maekawa; V. dentariifolia H. Boissieu; V. dissecta grasslands on mountain slopes; below 1300 m. Hebei, Jilin, Liaoning, Ledebour var. chaerophylloides (Regel) Makino; V. napellifolia Nei Mongol, Shaanxi, Shanxi [Mongolia, Russia]. Nakai; V. sieboldiana var. chaerophylloides (Regel) Nakai. One of us (Nikitin) prefers to treat Viola incisa as a distinct spe- cies, at least in Russia, and considers V. fissifolia also possibly to be a Terminal lobes highly variable in shape and size, ovate- separate species. lanceolate, lanceolate, oblong, or linear-lanceolate, margin ir- regularly dentate or shallowly divided, apex obtuse or acute. Fl. 45. Viola chaerophylloides (Regel) W. Becker, Bull. Herb. Mar–May, fr. May–Sep. 2n = 24*. Boissier, sér. 2, 2: 856. 1902. Mixed forests, forest margins, shaded and moist places in stream 南山堇菜 nan shan jin cai valleys, thickets on sunny slopes, grassy slopes; below 2000 m. Anhui, Chongqing, Hebei, Henan, Hubei, Jiangsu, Jiangxi, Liaoning, Shan- Herbs perennial, acaulescent, dwarf at anthesis, 4–20 cm dong, Zhejiang [Japan, Korea, SE Russia]. tall, to more than 30 cm tall at fruiting. Rhizome erect, short, 3– 10 mm, stout, surrounded by remains of stipules, with 3–6 ro- 45b. Viola chaerophylloides var. sieboldiana (Maximowicz) bust yellowish or white roots. Basal leaves 2–6, long petiolate; Makino, Bot. Mag. (Tokyo) 19: 87. 1905. stipules over 1/2 adnate to petioles, broadly lanceolate, mem- 细裂堇菜 xi lie jin cai branous, margin sparsely denticulate and ciliate or entire, apex acuminate; petioles 3–9 cm at anthesis, usually green, some- Viola pinnata Linnaeus var. sieboldiana Maximowicz, times purplish, glabrous, nitid, elongated at fruiting to more Bull. Acad. Imp. Sci. Saint-Pétersbourg 23: 313. 1877; V. than 20 cm; leaf blade glabrous on both surfaces or pubescent albida f. sieboldiana (Maximowicz) F. Maekawa; V. chaero- along veins, 3-sect, segments shortly stipitate, lateral segments phylloides f. sieboldiana (Maximowicz) F. Maekawa; V. dis- deeply 2-fid, central segment deeply 2- or 3-fid, terminal lobes secta f. sieboldiana (Maximowicz) Makino; V. dissecta var. highly variable in shape and size, ovate-lanceolate, lanceolate, sieboldiana (Maximowicz) Nakai; V. napellifolia Nakai var. oblong, or linear-lanceolate, margin irregularly dentate or shal- sieboldiana (Maximowicz) Naka; V. sieboldiana (Maxim- lowly divided, sometimes triparted to pinnatiparted or 3-lobed, owicz) Makino. apex obtuse or acute. Flowers white, creamy-white, or purplish, Terminal lobe of leaves triparted to pinnatiparted or 3- large, 2–2.5 cm in diam., fragrant; pedicels usually purplish, lobed. Fl. Mar–May, fr. May–Sep. nitid, subequaling or exceeding leaves at anthesis, glabrous, 2- bracteolate above middle; bracteoles linear or linear-lanceolate, Mixed forests, shaded and moist places in stream valleys, thickets margin very remotely and minutely denticulate. Sepals oblong- on sunny slopes, grassy slopes; below 1700 m. Anhui, Hubei, Jiangxi, Zhejiang [Japan]. ovate or narrowly ovate, 1–1.4 cm, basal auricles developed, 4.5–6 mm, glabrous, 3-veined, margin membranous, apex irreg- 46. Viola albida Palibin, Trudy Imp. S.-Peterburgsk. Bot. Sada ularly emarginate or shallowly divided. Petals broadly obovate, 17(1): 30. 1899. upper petals 1.3–1.5 cm × ca. 9 mm, lateral ones ca. 1.5 × 0.7 朝鲜堇菜 cm, sparsely bearded, anterior one purple-striate, 1.6–2 cm chao xian jin cai (spur included); spur long, 5–7 mm, stout, straight or slightly Herbs perennial, acaulescent, 5–20 cm tall, glabrous. Rhi- curved downward. Anthers 2.5–3 mm, spur of anterior 2 sta- zome erect, short, ca. 1 cm, densely noded, ca. 6 mm in diam., mens ca. 5 mm, slender, ca. 1.5 mm in diam. Ovary ca. 2 mm, with dense brown remains of stipules, with several yellowish glabrous; styles ca. 3 mm, base slightly geniculate; stigmas rootlets. Roots light colored, to 10 cm. Leaves basal, usually 2– slightly thickly margined on lateral sides and abaxially, slightly 4; stipules greenish, 1/3–1/2 adnate to petioles, free part linear- raised in central part, shortly beaked in front, with a rounded lanceolate, margin remotely denticulate; petiole usually longer

94 VIOLACEAE

than blades, sometimes shorter than blades at fruiting, slender, lanceolate, margin remotely and large toothed, apex acute. very narrowly winged in upper part, glabrous; leaf blade ovate Stipules ca. 1/2 adnate to petioles. Flowers white or light violet. or broadly ovate, entire or irregularly few lobed or 3–5-fid, 2– Fl. May, fr. Jun–Oct. 2n = 24. 11 × 1.5–7 cm, both surfaces glabrous, base deeply or shallowly Mixed forests; 300–500 m. Heilongjiang, Liaoning, Shandong cordate, margin regularly obtusely serrate, teeth curved toward [Korea]. inside, apex acuminate or shortly acuminate. Flowers white or light violet, purple striate, large, 1.6–2 mm in diam.; pedicels 1– In FRPS (51: 79. 1991), the name Viola savatieri Makino was 3, equaling or slightly exceeding leaves, slender, 2-bracteolate misapplied to this taxon. at or below middle; bracteoles linear, ca. 1.5 cm. Sepals oblong- 47. Viola forrestiana W. Becker, Repert. Spec. Nov. Regni lanceolate, ca. 1.6 cm (basal auricles included), apex acuminate, Veg. 19: 234. 1923. basal auricles conspicuous, 2.5–4.5 mm, margin narrowly membranous, apex irregularly 2- or 3-dentate. Petals obovate, 羽裂堇菜 yu lie jin cai ca. 1.6 cm, upper ones ca. 9 mm wide, lateral ones ca. 6 mm Herbs perennial, acaulescent, 5–12 cm tall. Rhizome erect, wide, sparsely bearded, anterior one ca. 2.2 cm (spur included), short; roots blackish, long, slender, smooth. Leaves basal, nu- apex emarginate; spur long, ca. 6 mm, stout, 3–4 mm in diam., merous; stipules mostly adnate to petioles, slightly white, lan- apex rounded. Anthers ca. 2 mm, appendage of connectives ca. ceolate, conspicuous, ca. 1.5 cm, membranous; petiole 2–7 cm, 2 mm; spur of 2 anterior stamens long, ca. 4 mm, slender, ca. glabrous; leaf blade triangular-ovate or narrowly ovate, 3–5.5 × 0.8 mm in diam. Ovary ca. 3 mm, glabrous; styles clavate, ca. 2–3.5 cm, abaxially glabrous, adaxially puberulous, base shal- 2.5 mm, base slender and slightly geniculate forward, gradually lowly cordate or sometimes subtruncate, margin irregularly thickened upward; stigmas slightly thickened and narrowly emarginate-crenate or acutely lobed below middle, lobes ob- margined on lateral sides and abaxially, slightly raised in cen- long and crenate, apex acute. Flowers purple or purplish; pedi- tral part and elongated forward into a short beak, with a cels usually exceeding or subequaling leaves, sparsely puberu- medium-sized stigma hole at tip of beak. Capsule ovoid-orbicu- lous, 2-bracteolate near middle; bracteoles linear-lanceolate, 5– lar, ca. 9 mm. 8 mm. Sepals lanceolate, ca. 5 mm, basal auricles ca. 2 mm, Mixed forests, thickets; 300–800 m. Heilongjiang, Liaoning, margin membranous, apex acute. Petals oblong-obovate, upper Shandong [Japan, Korea]. ones 1.2–1.4 cm, lateral ones subequaling upper ones, distinctly bearded, spur of anterior petal large, to 6 mm, stout, to 4 mm in 1a. Leaf blade entire ...... 46a. var. albida diam., slightly curved, apex thickened. Ovary glabrous; styles 1b. Leaf blade divided ...... 46b. var. takahashii clavate, slightly thickened upward; stigmas narrowly margined 46a. Viola albida var. albida on lateral sides and abaxially, flat in central part, shortly beaked in front, beak subhorizontal. Capsule globose. Fl. Mar–May, fr. 朝鲜堇菜 原变种 ( ) chao xian jin cai (yuan bian zhong) Jun–Sep.

Viola prionantha Bunge var. incisa Kitagawa, Rep. Inst. ● Grasslands on mountain slopes, streamsides, riversides; 2200– Sci. Res. Manchoukuo 6(4): 123. 1942; V. dissecta Ledebour 4000 m. Sichuan, SE Xizang. subvar. albida (Palibin) Makino; V. dissecta var. albida (Pali- bin) Nakai. 48. Viola gmeliniana Roemer & Schultes, Syst. Veg. 5: 354. 1819. Leaf blade ovate or broadly ovate, 2–11 × 1.5–7 cm, both 兴安堇菜 surfaces glabrous, base deeply or shallowly cordate, margin xing an jin cai regularly obtusely serrate, teeth curved toward inside, apex acu- Viola fusiformis Smith; V. gmeliniana var. albiflora W. minate or shortly acuminate; stipules ca. 1/3 adnate to petioles; Becker; V. gmeliniana var. glabra Ledebour; V. gmeliniana var. flowers white, sometimes reddish inside. Fl. Apr–May, fr. Jun– hispida Ledebour. Oct. 2n = 24. Herbs perennial, dwarf, acaulescent, 4–10 cm tall. Rhi- Mixed forests, thickets; 300–800 m. SE Liaoning [Japan, Korea]. zome erect, brown, thick, surrounded by remains of leaves of 46b. Viola albida var. takahashii (Nakai) Nakai, Bot. Mag. previous year in upper part. Roots usually blackish, smooth, (Tokyo) 36: 84. 1922. rarely branched. Leaves numerous, basal, rosulate; stipules greenish or glaucous, 1/2–3/4 adnate to petioles, the free part 菊叶堇菜 ju ye jin cai lanceolate, apex acute, margin remotely denticulate or sub- Viola dissecta var. takahashii Nakai, Icon. Pl. Koisik. entire; petiole inconspicuous or nearly absent on young leaves, 1(4): 94. 1912; V. albida f. takahashii (Nakai) Kitagawa; V. to 4 cm at fruiting, narrowly winged in upper part, stiffly hairy savatieri Makino f. detonsa (Kitagawa) Kitagawa; V. savatieri or subglabrous; leaf blade spatulate, long elliptic or lanceolate, var. detonsa Kitagawa; V. ×takahashii (Nakai) Takenouchi. 2–5 × 0.5–1.2 cm, subleathery, both surfaces ciliate or glabrous or densely stiffly hairy, or stiffly hairy only along veins adaxi- Leaf blade abaxially greenish, adaxially deep green, ovate ally, base narrow and decurrent to petiole, margin obtusely or oblong-ovate, 3–5 × 1.5–3 cm, glabrous, sometimes sparsely crenate or subentire and slightly repand, apex obtuse or ± acute. puberulous only along veins, base shallowly cordate, margin Flowers purple-violet, small; pedicels often purplish, usually greatly varied in appearance, usually irregularly few lobed or 3– exceeding or subequaling leaves, glabrous, rarely sparsely pu- 5-fid, lateral lobes shorter, middle lobe longer, terminal lobes berulous, 2-bracteolate at middle; bracteoles linear, 4–6 mm.

VIOLACEAE 95

Sepals ovate-lanceolate or lanceolate, 5–6 mm, margin nar- broad, 3-veined, margin narrowly membranous, apex rounded rowly membranous, apex acute, basal auricles subsquare, 1.5–2 or truncate, usually not dentate. Upper petals obovate, 1.1–1.3 mm, apex rounded or truncate, sometimes remotely dentate. cm × 5–8 mm, lateral ones oblong-obovate, 1.1–1.5 cm × 4–6 Petals oblong-obovate, upper ones ca. 9 × 5 mm, lateral ones mm, distinctly or sparsely bearded, anterior one 1.5–2.3 cm ca. 9 × 4.5 mm, distinctly bearded, anterior one 1–1.3 cm (spur (spur included); spur cylindric, long, 5–10 mm, robust, apex included); spur 2–3.5 mm, ca. 1.5 mm in diam., apex rounded, rounded, curved upward or straight. Appendage of connectives slightly curved upward. Anthers ca. 2 mm, appendage of con- ca. 1.5 mm; anthers ca. 2 mm; spur of 2 anterior stamens 4–6 nectives ca. 1.5 mm; spur of 2 anterior stamens short, ca. 3 mm, mm. Ovary ovoid-globose, ca. 2.5 mm, glabrous; styles clavate, stout, apex attenuate. Ovary ovoid-globose, ca. 1.8 mm, with base slender and geniculate forward, thicker in upper part; stig- granular appendages along abaxial ridge; styles ca. 2 mm, mas slightly thickened on lateral sides and abaxially and with clavate, base conspicuously geniculate forward, gradually thick- erect, thin margins, shortly beaked in front, beak obliquely as- ened upward; stigmas slightly thickened and margined on lat- cending, with a larger stigma hole at tip. Capsule oblong, 1–1.5 eral sides and abaxially, slightly raised at top, conspicuously cm, glabrous, apex acute. Seeds brownish red, numerous, ovoid- beaked in front, with a larger stigma hole at tip of beak. Capsule globose, ca. 1.5 mm, glabrous. Fl. Apr–May, fr. May–Sep. 2n = yellow-brown, narrowly ellipsoid, ca. 9 mm, glabrous. Fl. May– 24, 48*. Jun, fr. Jun–Aug. 2n = 24. Grasslands, grassy slopes, thickets, forest margins, sparse forests, Thickets on mountain slopes, river shores, sandy places, grassy fields, sandy places along river banks; below 1000 m. Anhui, Fujian, places on sand dunes; 300–1500 m. Heilongjiang, Nei Mongol [Mon- Heilongjiang, Jilin, Liaoning, Nei Mongol, Shandong, Taiwan [Japan, golia, Russia]. Korea, Russia]. 49. Viola mandshurica W. Becker, Bot. Jahrb. Syst. 54, Beibl. 50. Viola patrinii Candolle ex Gingins in Candolle, Prodr. 1: 120: 179. 1917. 293. 1824. 东北堇菜 dong bei jin cai 白花地丁 bai hua di ding Viola alisoviana Kiss f. intermedia (Kitagawa) Take- Viola patrinii var. brevicalcarata Skvortsov; V. primuli- nouchi; V. hsinganensis Takenouchi; V. mandshurica f. albi- folia var. glabra Nakai. flora P. Y. Fu & Y. C. Teng; V. mandshurica f. ciliata (Nakai) Herbs perennial, acaulescent, 7–20 cm tall. Rhizome erect, F. Maekawa; V. mandshurica var. ciliata Nakai; V. mand- dark brown, short, 4–10 mm, stout. Roots usually straight des- shurica f. glabra (Nakai) Hiyama ex Maekawa; V. mandshurica cending or slightly horizontal, black-brown, long, robust, often var. glabra Nakai; V. mandshurica f. macrantha (Maximowicz) produced from same place on rhizome. Leaves 3–5 or more, Nakai & Kitagawa; V. oblongosagittata Nakai f. ishizakii basal; stipules green, ca. 2/3 adnate to petioles, free part linear- Yamamoto; V. patrinii Candolle ex Gingins f. glabra (Nakai) F. lanceolate, margin remotely denticulate or entire, apex acumi- Maekawa; V. patrinii var. macrantha Maximowicz; V. philip- nate; petiole long, usually 2–3 × exceeding blades, 2–12 cm, pica Cavanilles f. intermedia (Kitagawa) Kitagawa; V. rhodo- slender, usually glabrous or sparsely puberulous, narrowly or sepala Kitagawa; V. yedoensis Makino f. intermedia Kitagawa. slightly broadly winged in upper part; leaf blade oblong, ellip- tic, narrowly ovate, or oblong-lanceolate, 1.5–6 × 0.6–2 cm, Herbs perennial, acaulescent, 6–18 cm tall. Rhizome erect, thin, both surfaces glabrous or puberulous along veins, base dark brown, short, 5–12 mm, densely noded, often producing truncate, slightly cordate, or broadly cuneate, decurrent to peti- from a single point several robust brown long roots; roots ole, margin remotely and shallowly repand-crenate or some- obliquely descending or sometimes slightly horizontal, usually times subentire, apex obtuse. Flowers white, purplish veined, smooth. Leaves 3–5 or numerous, basal; stipules membranous, medium-sized; pedicels usually exceeding or subequaling lower ones scalelike, brown, upper ones brownish, purplish, or leaves, slender, glabrous or sparsely puberulous, 2-bracteolate glaucous, over 2/3 adnate to petiole, free part linear-lanceolate, below middle; bracteoles linear. Sepals ovate-lanceolate or lan- apex acuminate, margin remotely denticulate or subentire; peti- ceolate, apex ± acute or slightly obtuse, basal auricles short, ca. ole 2.5–8 cm, narrowly winged in upper part, wing conspicu- 1 mm, obtuse. Upper petals obovate, ca. 1.2 cm, base narrow, ously broadened after anthesis, puberulous or glabrous; leaf lateral ones oblong-obovate, ca. 1.2 cm, bearded, anterior one blade oblong, ligulate, or ovate-lanceolate, lower ones usually ca. 1.3 cm (spur included); spur shallowly saccate, short, ro- smaller and narrowly ovate, 2–6 × 0.5–1.5 cm, accrescent after bust, ca. 3 mm or slightly shorter, ca. 3 mm in diam., apex anthesis, narrowly triangular, elliptic-lanceolate, or slightly has- rounded. Anthers ca. 2 mm, appendage of connectives ca. 1.5 tate, to 10 × 5 cm, broadest in lowest part, both surfaces gla- mm; spur of 2 anterior stamens short, ca. 2 mm, robust, ca. 0.6 brous or sparsely puberulous, midvein conspicuously raised mm in diam. Ovary narrowly ovoid, glabrous; styles clavate, abaxially, base truncate or broadly cuneate, decurrent to peti- slender, base slightly geniculate, slightly thickened in upper ole, margin remotely and shallowly repand-crenate or some- part; stigmas flat at top and triangular, narrowly margined on times subentire in lower part, apex obtuse or rounded. Flow- lateral sides, shortly beaked in front, beak obliquely ascending, ers purple-violet or purplish, sometimes white, ca. 2 cm in with a smaller stigma hole at tip. Capsule ca. 1 cm, glabrous. diam.; pedicels long, usually exceeding leaves, slender, gla- Seeds yellow-brown to dark brown, ovoid-globose. Fl. May– brous or puberulous, usually 2-bracteolate below or near mid- Jun, fr. Jun–Sep. 2n = 12, 24, 36. dle; bracteoles linear. Sepals ovate-lanceolate or lanceolate, 5–7 mm, apex acuminate, basal auricles short, 1.5–2 mm, Marshy meadows, moist places along riversides, thickets, moist

96 VIOLACEAE

and shaded places at forest margins; 200–1700 m. Heilongjiang, Jilin, light colored in lower part, 1.2–2.2 cm; pedicels usually pur- Liaoning, Nei Mongol [Japan, Korea, Mongolia, Russia]. plish red, unequal in length, exceeding or slightly shorter than 51. Viola striatella H. Boissieu, Bull. Herb. Boissier, sér. 2, 1: leaves, shortly hairy or subglabrous, 2-bracteolate at middle; 1077. 1901. bracteoles linear. Sepals usually purplish, oblong-lanceolate or ovate-lanceolate, 5–6 mm, 3-veined, margin narrowly mem- 圆叶堇菜 yuan ye jin cai branous or ciliate, apex acute, basal auricles shorter, 1–1.5 mm, with short stiff hairs or glabrous adaxially, apex truncate or re- Viola bambusetorum Handel-Mazzetti; V. emeiensis C. J. motely and shallowly dentate. Petals obovate, 7–14 mm, lateral Wang; V. lianhuashanensis C. J. Wang & K. Sun; V. pseudo- ones sparsely bearded or glabrous, anterior one white and vio- bambusetorum Chang; V. schensiensis W. Becker. let-striate at base, 1.2–2.2 cm (spur included); spur cylindric, Herbs perennial, acaulescent, 5–7 cm tall. Rhizome nearly 3–8 mm, robust or slender, apex obtuse, straight or slightly erect, slightly robust, shortly and conspicuously noded, with curved upward. Anthers and appendages of connectives ca. 2 few brown remains of stipules. Leaves basal, usually 3–5; mm; spur of 2 anterior stamens long, to 4 mm, slender, ca. 0.3 stipules brownish or dark brownish, adnate to petioles in lower mm in diam. Ovary subglobose, usually with short stiff hairs or part, free part ovate-lanceolate, margin entire or sparsely shortly subglabrous; styles clavate, base slightly geniculate, gradually fimbriate, apex acuminate; petiole long, 4–6 cm, slender; leaf thickened upward; stigmas conspicuously thickened on lateral blade subreniform, orbicular, or ovate, 1–1.5 × 1–1.5 cm, both sides and abaxially, shortly beaked in front, with a stigma hole surfaces glabrous, base narrowly cordate, margin obtusely ser- open upward at tip of beak. Capsule ellipsoid, ca. 7 mm, gla- rate, apex mucronate or obtuse. Flowers pinkish, deep purple, brous or sparsely short hairy, globose and usually with short purple-white, or white, small; pedicels slightly exceeding leaves, stiff hairs when young. Seeds brownish, small, ca. 1.5 mm, slender, 2-bracteolate near middle; bracteoles opposite, linear, shortly appendaged. Fl. Apr–May, fr. Jun–Sep. 2n = 24. ca. 5 mm, margin remotely denticulate. Sepals linear-lanceo- Grasslands on mountain slopes, forests, thickets, rock crevices in late, ca. 3 mm, margin narrowly membranous, apex ± acute, shaded places; 300–1700 m. Hebei, Heilongjiang, Jilin, Liaoning, Nei basal auricles not conspicuous, ca. 0.5 mm, apex obtuse. Petals Mongol, Shanxi [Japan, Korea, Mongolia, Russia]. oblong-obovate, upper ones ca. 8 × 3 mm, base attenuate, apex 53. Viola tenuicornis W. Becker, Beih. Bot. Centralbl., Abt. 2, obtuse, lateral ones ca. 7 × 3 mm, shortly bearded, anterior one 34: 248. 1916. ca. 8.5 mm (spur included); spur shortly saccate, 2–2.5 mm, ca. 细距堇菜 2 mm in diam., apex obtuse. Anthers ca. 1.5 mm, appendage of xi ju jin cai connectives subequaling anthers; spur of 2 anterior stamens Herbs perennial, acaulescent, 2–13 cm tall. Rhizome usu- shortly angular, ca. 1.5 mm. Ovary ovoid-globose, ca. 1.5 mm; ally erect, short, 2–10 mm, slender or slightly thick, densely styles clavate, 1.7–3 mm, base slender and slightly geniculate noded, with several yellowish rootlets. Leaves 2 to numerous, forward; stigmas slightly flat at top, narrowly margined on basal; outer stipules submembranous, inner ones greenish, 2/3 lateral sides, shortly beaked in front, beak erect, with a smaller adnate to petioles, free part linear-lanceolate or lanceolate, mar- stigma hole at tip. Fl. May, fr. Jun–Sept. 2n = 24*. gin remotely fimbriate-dentate; petiole 1.5–6 cm, slender, wing- ● Grasslands on mountain slopes, moist places on rocks; 1200– less or very narrowly winged in upper part, usually puberulous 3400 m. Anhui, Chongqing, Gansu, Henan, Hubei, Hunan, Jiangxi, or subglabrous; leaf blade abaxially dark purple to green, adax- Shaanxi, Sichuan, Yunnan. ially green, ovate or broadly ovate, 1–3 × 1–2 cm, accrescent at fruiting, to 6 × ca. 4.5 cm, glabrous or puberulous along veins 52. Viola variegata Fischer ex Link, Enum. Hort. Berol. Alt. 1: or margin, base slightly cordate or subrounded, margin shal- 240. 1821. lowly crenate, apex obtuse. Flowers purple-violet; pedicels 斑叶堇菜 ban ye jin cai slightly or not exceeding leaves, slender, puberulous or sub- glabrous, 2-bracteolate at or slightly below middle; bracteoles Viola tenuicornis W. Becker subsp. primorskajensis W. linear. Sepals usually green or purplish red, lanceolate or ovate- Becker; V. variegata f. viridis (Kitagawa) P. Y. Fu & Y. C. lanceolate, 5–8 × 1.5–2 mm, glabrous or puberulous, 3-veined, Teng; V. variegata var. viridis Kitagawa. margin narrowly membranous, apex acute, basal auricles short, Herbs perennial, acaulescent, 3–12 cm tall. Rhizome usu- 1–3 mm, apex truncate or rounded, rarely shallowly dentate. ally short, 4–15 mm, slender, densely noded, with several Petals obovate, upper ones 1–1.2 × ca. 6 mm, lateral ones 8–11 whitish or brownish long roots. Leaves basal, rosulate; stipules × 3–4.5 mm, slightly bearded or glabrous, anterior one 1.5– greenish or glaucous, submembranous, 2/3 adnate to petioles, 1.7(–2) cm (spur included); spur cylindric, 5–7(–9) mm, robust free part lanceolate, margin remotely glandular fimbriate-den- or slender, 1.2–3 mm in diam., apex rounded and curved up- tate, apex acuminate; petioles unequal in length, 1–7 cm, nar- ward. Anthers 1.5–2 mm; spur of 2 anterior stamens long, ca. rowly winged or wingless in upper part, with short stiff hairs or 5.5 mm, slender, ca. 0.3 mm in diam., apex rounded and slight- glabrous; leaf blade abaxially usually purplish red, adaxially ly curved. Ovary glabrous or puberulous; styles clavate, base dark green or green, white variegated (punctate-striate) along geniculate forward, conspicuously thickened in upper part; stig- veins, orbicular, 1.2–5 × 1–4.5 cm, both surfaces often covered mas conspicuously thickened on lateral sides and abaxially, with short stiff hairs, sometimes sparsely hairy or subglabrous, slightly raised in central part, shortly beaked in front, with a base conspicuously cordate, margin obtusely dentate, apex stigma hole open upward at tip of beak. Capsule ellipsoid, 4–6 rounded or obtuse. Flowers red-purple or dark purple, usually mm, glabrous.

VIOLACEAE 97

Forests, forest margins, thickets, mountain slopes, dry places in mm, base narrow, apex obtuse, lateral ones glabrous, anterior open fields; below 2300 m. Gansu, Hebei, Heilongjiang, Henan, Jiang- one slightly shorter, white veined; spur saccate, very short, su, Jilin, Liaoning, Nei Mongol, Shaanxi, Shandong, Shanxi [Korea, slightly longer than basal auricles of sepals or not. Anthers ca. Russia]. 1.5 mm, connective appendages ca. 1 mm, spur of 2 anterior sta- 1a. Leaf blade glabrous or puberulous mens very short, only ca. 0.4 mm. Ovary ovoid-globose, ca. 1.5 along veins or margin; sepals and mm, smooth, glabrous; styles clavate, base conspicuously gen- ovary glabrous ...... 53a. subsp. tenuicornis iculate, apex obtuse, very shortly beaked in front, with a smaller 1b. Leaf blade puberulous and verrucose; stigma hole at tip of beak. Capsule ovoid-orbicular, 5–7 mm. sepals white puberulous along Fl. May–Jul, fr. Jul–Aug. 2n = 20*. margin; ovary puberulous ...... 53b. subsp. trichosepala Alpine or subalpine meadows, subalpine thickets; 2900–4800 m. Gansu, Qinghai, NW Sichuan, Xinjiang, Xizang [Afghanistan, N India, 53a. Viola tenuicornis subsp. tenuicornis Kashmir, Kazakhstan, Kyrgyzstan, W Mongolia, Nepal, Russia, Tajiki- 细距堇菜(原亚种) xi ju jin cai (yuan ya zhong) stan]. Viola ×interposita Kitagawa; V. tenuicornis var. brach- 55. Viola magnifica C. J. Wang ex X. D. Wang, Acta Bot. ytricha W. Becker; V. variegata Fischer ex Link var. chinensis Yunnan. 13: 263. 1991. Bunge ex Regel; V. variegata var. viridis Kitagawa. 犁头叶堇菜 li tou ye jin cai Leaf blade glabrous or puberulous along veins or margin. Herbs perennial, acaulescent, ca. 28 cm tall. Rhizome Sepals and ovary glabrous. Fl. Apr–May, fr. Jun–Sep. 2n = 24*. robust, 1–2.5 cm, to 5 mm in diam., producing numerous terete Forests, forest margins, thickets, moist places in grasslands on roots and fibrous rootlets. Leaves basal, usually 5–7; stipules mountain slopes; 200–2300 m. Gansu, Hebei, Heilongjiang, Henan, large, 1/2–2/3 adnate to petioles, free part linear or narrowly Jiangsu, Jilin, Liaoning, Nei Mongol, Shaanxi, Shandong, Shanxi [Korea, lanceolate, ca. 1.2 cm, margin subentire or remotely denticulate; Russia]. petiole to 20 cm, very narrowly winged in upper part, glabrous; 53b. Viola tenuicornis subsp. trichosepala W. Becker, Beih. leaf blade adaxially deep green, triangular, triangular-ovate or Bot. Centralbl., Abt. 2, 34: 249. 1916. narrowly ovate, 7–15 × 4–8 cm, broadest at base, larger at fruit- ing, both surfaces glabrous, base broadly or deeply cordate, 毛萼堇菜 mao e jin cai lateral lobes spreading and large, margin distinctly coarsely Viola trichosepala (W. Becker) Juzepczuk. serrate, teeth obtuse and slightly incurved at apices, apex acu- minate. Flowers large, light pink or bluish purple. Sepals nar- Petiole appressed puberulous; leaf blade puberulous and rowly ovate, 4–7 × 2–3.2 mm; basal auricles 3–5 mm, apex verrucose. Sepals white puberulous along margin. Lateral petals deeply dentate. Lateral petals slightly bearded, anterior petal distinctly bearded. Ovary puberulous. oblong, ca. 1.9 cm (spur included); spur 4–5 mm, 3–3.5 mm in Sunny mountain slopes, dry places in open fields; below 1900 m. diam. Capsule ellipsoid, 1.2–2 cm, ca. 5 mm in diam., glabrous; Hebei, Jilin, Liaoning, Nei Mongol, Shanxi [Korea, Russia]. fruit stalk 4–15 cm, glabrous, 2-bracteolate near or below mid- 54. Viola kunawarensis Royle, Ill. Bot. Himal. Mts. 75. t. 18. dle; bracteoles linear or linear-lanceolate, 7–10 mm. Fl. Mar– 1839. Apr, fr. Jul–Oct. 西藏堇菜 xi zang jin cai ● Forests on mountain slopes, shaded and moist places at forest margins or in valleys; 800–2000 m. Anhui, Chongqing, NW Guizhou, Viola kunawarensis var. angustifolia W. Becker; V. kuna- Henan, Hubei, Hunan, Jiangxi, Zhejiang. warensis f. longifolia C. J. Wang; V. thianschanica Maxim- 56. Viola sphaerocarpa W. Becker, Beih. Bot. Centralbl., Abt. owicz. 2, 36: 54. 1918. Herbs perennial, dwarf, acaulescent, 2.5–6 cm tall. Rhi- 圆果堇菜 zome short, robust, densely noded; roots brownish or glaucous, yuan guo jin cai conic, long, slender, usually simple. Leaves basal, rosulate; stip- Viola keiskei Miquel f. hirsutior W. Becker. ules whitish, 1/2–2/3 adnate to petioles, membranous, free part lanceolate, margin sparsely glandular fimbriate, apex acu- Herbs perennial, acaulescent, 6–12 cm tall, sparsely pu- minate; petiole slightly longer than or nearly as long as leaf berulous. Rhizome erect, whitish, slender, 2–2.5 mm in diam. blade; leaf blade ovate, elliptic, or oblong, 0.5–2 cm × 2–5 mm, inconspicuously noded, with branched rootlets in lower part. thickly papery, both surfaces glabrous, midvein conspicuously Leaves basal, numerous; stipules ca. 1/3 adnate to petioles at raised, base cuneate or broadly cuneate, slightly decurrent, mar- base, the free part subulate-linear, 0.8–1.4 cm, margin remotely gin entire or remotely and shallowly crenate, apex obtuse. Flow- glandular fimbriate-dentate; petiole to 13 cm, sparsely pubes- ers deep blue-purple, small; pedicels straight, slightly longer cent, narrowly winged in the upper part; leaf blade gray-green, than or nearly as long as leaves, slender, 2-bracteolate slightly orbicular-cordate or ovate-orbicular, 2.5–6.5 × 2–4.5 cm, both above middle; bracteoles nearly opposite, linear or narrowly lan- surfaces sparsely puberulous, more densely puberulous along ceolate, sparsely glandular-fimbriate along lower margin, apex veins abaxially, base deeply or broadly cordate, margin obtuse- acuminate. Sepals oblong or ovate-lanceolate, 3–4 × 1–1.5 mm, serrate, apex ± acute or acuminate. Pedicels 3–6 cm, glabrous, apex obtuse, basal auricles very short, 3-veined, margin nar- with 2 ciliate bracteoles about the middle. Sepals ovate-lanceo- rowly membranous, apex obtuse. Petals oblong-obovate, 7–10 late, puberulous along veins abaxially, basal auricles 2–3 mm,

98 VIOLACEAE

apex enlarged, dentate. Petals white, lateral petals bearded, an- Herbs perennial, acaulescent, 8–10 cm tall, gray-green, terior one 1.5–1.8 cm including spur, spur ca. 6 mm, ca. 2 mm glabrous. Rhizome erect, 7–8 cm, robust, 3–4 mm in diam., in diam. Ovary glabrous. Capsule globose, 6–10 mm, glabrous. without rootlets. Leaves basal, gray-green; stipules narrowly Seeds ca. 1.8 mm, ca. 1 mm in diam. Fl. Mar, fr. May–Oct. lanceolate, ca. 1 cm, ca. 2/3 adnate to petiole, margin sparsely shortly fimbriate; petiole subequaling leaf blade, narrowly ● Mountain forests, forest margins, thickets; 1200–3000 m. winged in upper part; leaf blade lanceolate, 3–4 × ca. 1 cm, Chongqing, S Shaanxi, Sichuan, NW Yunnan. both surfaces glabrous, veins raised abaxially, base narrowly 57. Viola inconspicua Blume, Bijdr. 58. 1825. cuneate, long decurrent to petiole, margin remotely and shal- lowly serrate, apex ± acute. Flowers purplish; pedicels usually 长萼堇菜 chang e jin cai exceeding leaves. Sepals ovate-lanceolate, ca. 5 mm, ca. 1 mm Viola betonicifolia Smith subsp. novaguineensis D. M. in diam., apex acuminate, basal auricles nearly square, nar- Moore; V. betonicifolia var. oblongosagittata (Nakai) F. rower. Petals elliptic, lateral ones bearded, anterior one spatu- Maekawa & T. Hashimoto; V. chinensis G. Don; V. confusa late, shortly spurred; spur ca. 2 mm, slightly robust. Ovary Champion ex Bentham; V. confusa subsp. nagasakiensis (W. glabrous; styles clavate, base slightly geniculate, gradually Becker) F. Maekawa & T. Hashimoto; V. hunanensis Handel- thickened in upper part; stigmas with erect margins on lateral Mazzetti; V. inconspicua subsp. nagasakiensis (W. Becker) J. sides and abaxially, shortly beaked in front, beak slightly erect. C. Wang & T. C. Huang; V. mandshurica subsp. nagasakiensis Capsule ellipsoid, 5–6 mm, glabrous. Fl. Apr and Sep, fr. May– Oct. W. Becker; V. minor (Makino) Makino; V. oblongosagittata Nakai; V. patrinii var. minor Makino; V. philippica subsp. ● Grasslands on mountain slopes; elevation unknown. W Hubei, malesica W. Becker; V. primulifolia Loureiro (1790), not Hunan. Linnaeus (1753); V. pseudomonbeigii Chang; V. serpens 59. Viola betonicifolia Smith in Rees, Cycl. 37: Viola no. 7. Wallich ex Gingins var. confusa (Bentham) J. D. Hooker & 1817. Thomson. 戟叶堇菜 ji ye jin cai Herbs perennial, acaulescent. Rhizome erect or obliquely ascending, 1–2 cm, robust, 2–8 mm in diam., densely noded, Viola betonicifolia subsp. dielsiana W. Becker; V. with white roots. Leaves basal, rosulate; stipules ca. 3/4 adnate betonicifolia subsp. nepalensis (Gingins) W. Becker; V. to petiole, free part lanceolate, 3–7 mm, margin sparsely shortly caespitosa D. Don; V. inconspicua Blume subsp. dielsiana W. fimbriate-dentate, rarely entire, usually brown rusty dotted, apex Becker; V. oblongosagittata Nakai var. violascens Nakai; V. acute; petiole 2–7 cm, usually glabrous or rarely shortly pubes- patrinii Gingins var. caespitosa (D. Don) Ridley; V. patrinii cent; leaf blade triangular-ovate, 1.5–9 × 1–7 cm, 1–2.2 × as var. laotiana H. Boissieu; V. patrinii var. nepaulensis Gingins. long as wide, broadest at base, usually glabrous, rarely puberu- Herbs perennial, acaulescent. Rhizome obliquely ascend- lous, base cordate with usually prominent, rounded basal lobes, ing or erect, usually short, 5–10 mm, stout, with several robust, decurrent to 1/3 length of petiole and narrowly winged, apex long, yellowish roots. Leaves numerous, basal, rosulate; stip- acuminate or acute. Flowers purplish or white, with darker stri- ules dark brownish, lanceolate, 3–5 × ca. 1 mm, ca. 3/4 adnate ations; pedicels usually subequaling or slightly exceeding leaves, to petiole, free part linear-lanceolate or subulate, margin entire slender, glabrous or puberulous in upper part, 2-bracteolate or sparsely denticulate, apex acuminate; petiole long, 1.5–13 slightly above middle; bracteoles linear. Sepals ovate-lanceo- cm, narrowly and conspicuously winged in upper part, usually late, 4–7 × 1–1.8 mm, apex acute, glabrous or ciliate; append- glabrous, sometimes puberulous in lower part; leaf blade nar- age elongated, (1.5–)2–3(–4.8) mm, shallowly incised. Petals rowly lanceolate, narrowly triangular-hastate, or triangular- oblong-obovate, 7–9 mm; lateral ones bearded or rarely gla- ovate, 2–7.5 × 0.5–3 cm, base truncate or shallowly cordate, brous; anterior one 1–1.2 cm (spur included); spur tubular, 1.8– sometimes broadly cuneate, apex acute, sometimes slightly ob- 3(–4) mm, straight, obtuse. Spur of anterior stamens angular, tuse; leaves accrescent after anthesis, both surfaces glabrous or base broad, apex acute. Ovary globose, glabrous; styles clavate, subglabrous, lateral lobes spreading and conspicuously dentate, base slightly geniculate; stigmas flat at top, with broader margin margin remotely shallowly crenate, more deeply dentate near on lateral sides, shortly beaked in front, with a stigmatic hole base. Flowers white, light or deep purplish, deep-colored striate, opening upward at tip of beak. Capsule oblong, 8–10 mm, 1.4–1.7 cm; pedicels long, equaling or exceeding leaves, slen- glabrous. Seeds deep green, ovoid-globose, 1–1.5 mm, ca. 0.8 der, usually glabrous, sometimes puberulous in lower part, 2- mm in diam. Fl. Nov–Apr, fr. Jun–Nov. 2n = 24*, 48*, 72. bracteolate near middle; bracteoles linear. Sepals ovate-lanceo- Grasslands, field sides, path margins, forest margins; below late or lanceolate, 5–6 × 1–1.6 mm, apex acuminate or ± acute, 1600(–2400) m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, basal auricles shorter, 0.5–1 mm, 3-veined, margin narrowly Henan, Hubei, Hunan, Jiangsu, Jiangxi, SE Shaanxi, Sichuan, Taiwan, membranous, apex rounded, sometimes remotely obtusely den- Yunnan, Zhejiang [India, Indonesia, Japan, Malaysia, Myanmar, New tate. Upper petals obovate, 1–1.2 cm, bearded; lateral ones ob- Guinea, Philippines, Vietnam]. long-obovate, bearded, 1–1.2 cm; anterior one usually shorter, 58. Viola cuspidifolia W. Becker, Bull. Misc. Inform. Kew 1.3–1.5 cm (spur included); spur tubular, short, 1–2.5 mm, 1929: 201. 1929. stout, 2–3 mm in diam., apex rounded, straight or slightly curved upward; anthers and appendage of connectives ca. 2 鄂西堇菜 e xi jin cai mm; spur of 2 anterior stamens 1–3 mm. Ovary ovoid-globose,

VIOLACEAE 99

ca. 2 mm, glabrous; styles clavate, slightly geniculate forward, to ascending, rather stout, with white or yellowish roots. Stip- gradually thickened in upper part; stigmas slightly thickened ules narrowly lanceolate, 1/3–1/2 adnate to petioles, margin en- and narrowly margined on lateral sides and abaxially, shortly tire or inconspicuously and remotely glandular serrate; petiole beaked in front, with a stigma hole at tip of beak. Capsule ellip- sparsely puberulous or glabrous, narrowly winged; leaf blade soid to oblong, 6–9 mm, glabrous. Seed ovoid, 1–1.2 mm. Fl. oblong-ovate, 3–8 × 1.5–5 cm, both surfaces appressed puberu- Feb–May, fr. May–Sep. 2n = 24*, 48, 72*. lous or glabrescent, base shallowly cordate or truncate and long decurrent to petiole, margin serrate to crenate-serrate, apex Fields, roadsides, grasslands on mountain slopes, thickets, forest slightly acute. Pedicels longer than leaves, 7–9 cm, glabrous or margins; below 1500(–2500) m. Anhui, Chongqing, Fujian, Guang- very sparsely puberulous, 2-bracteolate near middle; bracteoles dong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan, Jiangsu, Jiang- xi, Shaanxi, Sichuan, Taiwan, SE Xizang, Yunnan, Zhejiang [Afghani- linear, ca. 5 mm. Sepals ovate or ovate-lanceolate, ca. 5 × 1.8–2 stan, Bhutan, Japan, India, Indonesia, Kashmir, Malaysia, Myanmar, mm, appendages very narrowly square, 1–1.5 mm, apex obtuse Nepal, Philippines, Sri Lanka, Thailand, Vietnam; N Australia]. or dentate. Petals rounded at apex, upper ones obovate, ca. 9 × 4.5 mm, lateral ones oblong, ca. 1 × 0.3 cm, bearded; anterior 60. Viola trichopetala C. C. Chang, Bull. Fan Mem. Inst. Biol., one triangular-obovate, ca. 9 mm; spur somewhat elongated, 4– n.s., 1: 254. 1949. 6 mm, thin, somewhat curved upward. Style somewhat genicu- 毛瓣堇菜 mao ban jin cai late at base, clavate, apex flat, broadened on both sides and distinctly margined, somewhat erectly beaked in front. Capsule Viola betonicifolia Smith f. pubescens H. Hara; V. philip- ellipsoid, 5–7 mm, glabrous. Fl. Mar–May, fr. May–Oct. pica Cavanilles var. yunnanfuensis W. Becker. Grassy slopes, thickets or sparse forests; 300–3200 m. Guizhou, Herbs perennial, acaulescent, to 7 cm tall. Rhizome erect, Sichuan, SE Xizang, Yunnan [India, Kashmir, Nepal]. brownish, 1.5–2.5 cm, slightly robust; roots straightly descend- 62. Viola senzanensis Hayata, Icon. Pl. Formosan. 6: 3. 1916. ing, long, robust. Leaves usually numerous, basal; stipules brown, 1/2–2/3 united with petioles, free part linear-lanceolate, 尖山堇菜 jian shan jin cai margin entire or sparsely brown glandular denticulate, apically Herbs perennial. Rhizome long creeping to ascending, slen- acuminate; petioles exceeding or shorter than blades, narrowly der, acaulescent. Stipules lanceolate, 5–10 × 1–2 mm, glabrous, winged in upper part, with sparse spreading short stiff hairs or membranous, ca. 2/3 adnate to petiole, margin slightly lacini- glabrous; blades in upper part lanceolate, those in lower part ate-serrate or entire, apex acute; petiole 2–11 cm, glabrous; leaf triangular-ovate or oblong-ovate, 3–4.5 × 2–3 cm, both surfaces blade triangular-cordate, 1.2–2.8 × 2–3.8 cm, herbaceous to pubescent or nearly glabrous, base broadly cordate or cuneate, papery, glabrous or puberulous on both sides, base reniform- decurrent to petiole, margin sparsely obtuse-crenate, apex ob- cordate, with basal lobes somewhat converging, margin shal- tuse. Flowers purplish violet or white with purple veins, ca. 1.5 lowly serrate, wingless, apex acute to obtuse. Flowers nearly cm in diam.; pedicels exceeding leaves at anthesis, 2-brac- white, with dark violet veins, 1–1.5 cm in diam.; pedicels 4–10 teolate below middle; bracteoles linear, 5–7 mm, margin entire. cm, exceeding leaves, slender, glabrous or sparsely puberulous. Sepals lanceolate, ca. 5 mm, ca. 1.5 mm wide at base, apex Sepals ovate-lanceolate to lanceolate, 3.5–6 × 0.8–1.5 mm, gla- acute; basal auricles short, ca. 0.5 mm, glabrous, 3-veined, mar- brous, margin entire, apex acute; appendages ca. 1 mm, round- gin entire or sometimes repand-crenate, apex obtuse. Petals obo- ed. Petals obovate to oblong, lateral ones glabrous, anterior one vate, margin entire, lateral petals densely bearded, upper petals equaling others, apex rounded; spurs cylindric, 2–3 mm, slen- also few bearded, anterior petal ca. 1.5 cm (spur included); spur der, rounded. Styles 2–2.7 mm, base slightly geniculate, clavate shallowly saccate, short, 1–2(–3) mm, stout, ca. 2 mm thick, distally, apex slightly lobed on both sides; stigmas beaked. Cap- apex rounded. Ovary glabrous, oblong; styles slender, clavate, sules oblong or ellipsoid, 6–9 mm, glabrous. Seeds ovoid, ca. 2 geniculate at base, ca. 1.5 mm; stigmas with slightly thin and mm. 2n = 24*. straightly spreading margin on 2 lateral sides and behind, slightly convex in central part, shortly beaked in front, with ● High mountains; 3300–3600 m. Taiwan. smaller stigma hole at tip of beak. Capsule oblong, 1–1.2 cm, 63. Viola philippica Cavanilles, Icon. 6: 19. 1801. glabrous, apex acute. Seeds red-brown, ovoid-globose, ca. 2 mm, ca. 1.5 mm thick. Fl. Feb–Jun, fr. Jun–Dec. 紫花地丁 zi hua di ding Grassy slopes, thickets; 1600–3400 m. SW Sichuan, Xizang, Yun- Herbs perennial, acaulescent, 4–14 cm tall, to 20 cm tall at nan [Bhutan]. fruiting. Rhizome erect, brownish, short, 4–13 mm, 2–7 mm in diam., densely noded, with several yellowish or whitish root- 61. Viola tienschiensis W. Becker, Repert. Spec. Nov. Regni lets. Leaves numerous, basal, rosulate; stipules glaucous or Veg. 17: 314. 1921. greenish, 1.5–2.5 cm, membranous, 2/3–4/5 adnate to petioles, 滇西堇菜 dian xi jin cai free part linear-lanceolate, margin remotely glandular fimbriate- denticulate or subentire; petiole usually 1–2 × exceeding blades Viola angustistipulata C. C. Chang; V. betonicifolia Smith at anthesis, very narrowly winged in upper part, to 10 cm at subsp. jaunsariensis (W. Becker) H. Hara; V. ganchouenensis fruiting, broadly winged in upper part, puberulous or glabrous; W. Becker; V. flavida Bureau & Franchet (1891), not Schur upper blades oblong-lanceolate or triangular-ovate, usually (1877); V. oligoceps Chang; V. prionantha subsp. jaunsariensis smaller than lower ones; lower ones oblong, narrowly ovate- W. Becker. lanceolate, or oblong-ovate, 1.5–4 × 0.5–1 cm, both surfaces Herbs perennial, acaulescent, to 12 cm tall. Rhizome erect glabrous or puberulous, sometimes only puberulous along veins

100 VIOLACEAE

abaxially, base truncate or cuneate, rarely slightly cordate, mar- Basionym: Viola pseudojaponica Nakai, Bot. Mag. (To- gin shallowly crenate, apex obtuse; blades accrescent at fruit- kyo) 42: 560. 1928 [“pseudo-japonica”]; V. longistipulata Ha- ing, to 10 × 4 cm. Flowers purple-violet or purplish, rarely yata; V. nantouensis S. S. Ying; V. stenocentra Hayata ex white, light colored and purple-striate at throat, medium-sized; Nakai; V. taiwaniana Nakai; V. yedoensis var. pseudojaponica pedicels usually numerous, equaling or exceeding leaves, slen- (Nakai) T. Hashimoto ex E. Hama & K. Nakai. der, glabrous or puberulous, 2-bracteolate near middle; bracte- oles linear. Sepals ovate to ovate-lanceolate, 3–7 × 0.7–1.8 mm, Leaves present in winter; petiole distinctly winged; leaf apex acuminate; appendages short, 1–1.5 mm, 1/2 to 1/6 as long blade triangular-lanceolate, 3–7 × 3.5–5 cm, usually glabrous, as sepal, glabrous or sparsely pubescent, apex rounded or den- rarely pubescent, base truncate or slightly cordate, apex acu- ticulate. Petals obovate or oblong-obovate, lateral ones 1–1.2 minate. Fl. Nov–Apr, fr. May–Oct. 2n = 24*. cm, inside glabrous or rarely lightly bearded, anterior one 1.3–2 Roadsides. Taiwan [Japan (S Kyushu and Ryukyu Islands)]. cm (spur included), inside purple-veined; spur tubular, 3–8 mm, 2–5 × as long as calycine appendages; anthers ca. 2 mm, append- 64. Viola mongolica Franchet, Pl. David. 1: 42. 1884. age of connectives ca. 1.5 mm; spur of 2 anterior stamens 蒙古堇菜 meng gu jin cai tubular, 3–6 mm, apex slender. Ovary ovoid, glabrous; styles clavate, slightly exceeding ovary, base slightly geniculate; stig- Viola hebeiensis J. W. Wang & T. G. Ma; V. yamatsutae mas triangular with slightly thickened and raised margins on Ishidoya ex Kitagawa; V. yezoensis Maximowicz var. hebei- lateral sides and abaxially, slightly flat at top, shortly beaked in ensis (J. W. Wang & T. G. Ma) J. W. Wang & J. Yang front. Capsule ellipsoid, 5–12 mm, glabrous. Seeds yellowish, [“hopeiensis”]. ovoid-globose, ca. 1.8 mm. Fl. Apr–May, fr. May–Sep. 2n = Herbs perennial, acaulescent, to 15 cm tall. Rhizome erect 24*, 48. or obliquely ascending, 0.5–2 cm, robust, to 0.5 cm thick, with Fields, grassy places on mountain slopes, forest margins, thickets, numerous brownish robust roots. Leaves basal; stipules green- roadsides; below 1700 m. Anhui, Chongqing, Fujian, Gansu, Guang- ish, 1/2 adnate to petioles, free part lanceolate, margin sparsely dong, Guangxi, Guizhou, Hainan, Hebei, Heilongjiang, Henan, Hubei, and shortly fimbriate-dentate, apex acute; petiole 3–4 cm, to 12 Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shan- cm at fruiting, puberulous, narrowly winged; blade ovate or long dong, Shanxi, Sichuan, Taiwan, Yunnan, Zhejiang [Cambodia, India, Indonesia, Japan, Korea, Laos, Mongolia, Philippines, Vietnam]. ovate, 2–5 × 3–4 cm, to 8 × 4.5 cm at fruiting, both surfaces puberulous, base deeply cordate, sometimes shallowly cordate, 1a. Leaves withered in winter, blade oblong- margin shallowly serrate, apex acute or obtuse. Flowers white, lanceolate to triangular-lanceolate, throat yellowish green, long pedicellate; pedicels robust, usu- petioles slightly winged ...... 63a. var. philippica ally exceeding leaves, 6–8 cm, puberulous, 2-bracteolate in or 1b. Leaves present in winter, blade above middle; bracteoles linear, 1–1.5 cm, margin sparsely triangular-lanceolate, petioles pilose. Sepals lanceolate, usually pilose, 1.1–1.3 cm (basal auri- rather distinctly winged ...... 63b. var. pseudojaponica cles included), 3–4 mm wide, apex acute; basal auricles con- 63a. Viola philippica var. philippica spicuous, 3–4 mm, apex deeply or shallowly emarginate. Upper petals obovate, ca. 1.2 × 0.8 cm, base narrowed and clawed, lat- 紫花地丁(原变种) zi hua di ding (yuan bian zhong) eral ones oblong-obovate, ca. 1.3 × 0.6 cm, sparsely bearded and yellowish at throat, anterior one 1.8–2 cm (spur included), Viola alisoviana Kiss; V. alisoviana f. candida (Kitagawa) with 5–7 dark veins, apex obtuse; spur cylindric, robust, 5–7 Takenouchi; V. chinensis G. Don f. alboviolacea Skvortsov; V. chinensis f. anomala Skvortsov; V. chinensis f. communis mm, ca. 2.5 mm thick, apex obtuse, usually curved upward or Skvortsov; V. chinensis f. dissecta Skvortsov; V. chinensis f. straight. Anthers subequaling membranous apical appendages glabra Skvortsov; V. philippica f. candida (Kitagawa) Kita- of connectives, ca. 2 mm; spur of anterior stamens narrowly gawa; V. philippica subsp. munda W. Becker; V. patrinii Gin- linear, ca. 5 mm, usually curved near apex. Ovary glabrous; gins var. chinensis Gingins; V. yedoensis Makino; V. yedoensis styles usually erect at base, thicker in upper part; stigmas nar- f. candida Kitagawa. rowly margined on lateral sides and behind, shortly and thickly beaked in front, with a larger stigma hole at tip of beak. Capsule Plant glabrous or pubescent. Leaves withered in winter; oblong, ca. 1 cm. Fl. Apr–May, fr. May–Oct. petiole slightly winged; leaf blade oblong-lanceolate to triangu- lar-lanceolate. Fl. Apr–May, fr. May–Sep. 2n = 24*, 48. ● Broad-leaved forests, thickets on mountains, grasslands on mountain slopes; (200–)800–2800 m. Gansu, Hebei, Heilongjiang, Fields, grassy places on mountain slopes, forest margins, thickets; Henan, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shan- below 1700 m. Anhui, Chongqing, Fujian, Gansu, Guangdong, Guang- dong, Shanxi. xi, Guizhou, Hainan, Hebei, Heilongjiang, Henan, Hubei, Jiangsu, In FRPS (51: 44. 1991), the name Viola yezoensis Maximowicz Jiangxi, Jilin, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shandong, was misapplied to this species. Shanxi, Sichuan, Taiwan, Yunnan, Zhejiang [Cambodia, India, Indo- nesia, Japan, Korea, Laos, Mongolia, Philippines, Vietnam]. 65. Viola hancockii W. Becker, Bull. Misc. Inform. Kew 1928: 63b. Viola philippica var. pseudojaponica (Nakai) Y. S. Chen, 249. 1928. comb. nov. 西山堇菜 xi shan jin cai 琉球堇菜 liu qiu jin cai Viola hancockii var. fangshanensis J. W. Wang.

VIOLACEAE 101

Herbs perennial, acaulescent, 10–15 cm tall. Rhizome 1.5– upper part; stigmas conspicuously margined on lateral sides 2 cm, robust, 4–6 mm in diam., densely noded. Roots deep and abaxially, shortly beaked in front, with a larger stigma hole brown, long, robust, often branched, longest to 13 cm, with nu- at tip of beak. Capsule ellipsoid, ca. 1 cm, glabrous. Fl. Apr– merous rootlets. Leaves numerous, basal; outer stipules white, Jul, fr. Jul–Sep. 1–1.3 cm × ca. 4 mm, membranous, inner ones ca. 3/4 adnate to ● Forest margins, grassy places on shaded mountain slopes, petiole, free part broadly lanceolate or lanceolate, margin re- streamsides in mountain valleys; 1900–3200 m. Sichuan, SW Xizang, motely shortly dentate; petiole slender, wingless, equaling or Yunnan. slightly exceeding blades, sparsely puberulous or glabrous; leaf blade abaxially pink to pinkish green, adaxially green or con- 67. Viola pekinensis (Regel) W. Becker, Beih. Bot. Centralbl., spicuously white punctate-striate along veins, ovate-cordate, 2– Abt. 2, 34: 251. 1916. 6 × 2–4 cm, abaxially sparsely puberulous at base or subgla- 北京堇菜 bei jing jin cai brous, veins conspicuously raised, adaxially sparsely puberu- lous, base deeply cordate, sinus narrow or slightly spreading, Viola dolichoceras C. J. Wang; V. kamtschatica Gingins margin regularly obtusely serrate, apex acute, sometimes obtuse. var. pekinensis Regel; V. liaosiensis P. Y. Fu & Y. C. Teng; V. Flowers whitish, large, to 2 cm; pedicels usually not or some- mongolica f. longisepala P. F. Fu & Y. C. Teng. times slightly exceeding leaves, 2-bracteolate at middle; brac- Herbs perennial, acaulescent, 5–9 cm tall, to 17 cm at teoles alternate, linear, 8–10 mm, margin remotely denticulate, fruiting, usually with persistent remains of leaves of last year at apex acuminate. Sepals lanceolate or broadly lanceolate, 7–8 × anthesis. Rhizome erect or obliquely ascending, 1–4 cm or ca. 3 mm, apex acute, basal auricles short, ca. 1.5 mm, remotely longer, slightly robust, with numerous white rootlets. Leaves obtusely dentate, apex truncate. Petals oblong-obovate, upper several, basal; stipules 1/2–2/3 adnate to petioles, free part nar- ones ca. 1.2 × 0.8 cm, lateral ones bearded, ca. 1.2 × 1 cm, an- rowly lanceolate, margin remotely denticulate; petiole narrowly terior one 1.8–2 cm (spur included); spur cylindric, 6–8 mm, winged, 2–7 cm, glabrous; leaf blade ovate-cordate, cordate, or apex rounded, usually curved upward; spur of anterior stamens elliptic-cordate, 1.5–3 × 1–2 cm, larger at fruiting, 2.5–6 × 2–5 angular, 4–5 mm, slender. Ovary subglobose, glabrous; styles cm, both surfaces sparsely puberulous, abaxially sometimes slightly geniculate at base; stigmas flat at apex, with slightly subglabrous, base cordate, margin obtusely serrate, apex obtuse thick margin, shortly beaked in front, with a small stigma hole or acute. Flowers white or rarely light rose; pedicels usually at tip. Capsule oblong, 0.7–1 cm, glabrous. Fl. Apr–May, fr. exceeding leaves, slender, glabrous, 2-bracteolate near middle; Jun–Sep. 2n = 24*. bracteoles linear. Sepals elliptic-lanceolate or oblong-lanceo- ● Broad-leaved forests, forest margins, streamsides; 200–1800 m. late, 7–9 mm, apex obtuse or acute; basal auricles 1.5–2.5 mm, Gansu, Hebei, Henan, Jiangsu, Shaanxi, Shandong, Shanxi. apex truncate or shallowly dentate, ciliate. Lateral petals beard- ed, anterior one 1.5–2 cm (spur included), sometimes purple 66. Viola belophylla H. Boissieu in Bull. Soc. Bot. France 55: striate in middle and lower parts; spur slightly curved upward, 467. 1908. cylindric, 6–8 mm, apex obtuse. Ovary glabrous; styles slightly 枪叶堇菜 qiang ye jin cai geniculate forward at base, gradually thickened upward; stig- mas with wider margin on two lateral sides and behind, shortly Viola monbeigii W. Becker. beaked in front, with a slightly upward stigma hole at tip. Cap- sule ellipsoid or ovoid, 6–8 mm, glabrous. Fl. Apr–May, fr. Herbs perennial, acaulescent, 8–15 cm tall. Rhizome whit- Jun–Aug. ish to brown, ligneous, long, to 5.5 cm, 3–6 mm in diam., dense- ly noded, with several, scattered, white roots. Leaves basal; ● Broad-leaved forests, needle-leaved forests, forest margins, stipules whitish or green, outer ones ovate-lanceolate, margin stony places, crevices; 500–1900 m. Hebei, Heilongjiang, Henan, Jilin, entire, inner ones ca. 1/2 adnate to petiole, free part narrowly Liaoning, Nei Mongol, Shandong, Shanxi. lanceolate, margin remotely denticulate, apex acuminate; peti- In FRPS (51: 42. 1991), the name Viola mongolica was mis- ole 2.5–6(–20) cm, narrowly winged in upper part; leaf blade applied to this species. abaxially pale green, adaxially deep green, ovate or narrowly 68. Viola lactiflora Nakai, Bot. Mag. (Tokyo) 28: 329. 1914. ovate, 2–20 × 1.5–7 cm, both surfaces subglabrous or young leaves puberulous abaxially, base deeply cordate, sometimes 2 白花堇菜 bai hua jin cai lobes overlapping, margin obtusely serrate, apex acute or cau- Viola limprichtiana W. Becker. date-acuminate. Flowers white, 1.7–1.9 cm; pedicels purplish in lower part, usually exceeding or subequaling leaves at an- Herbs perennial, acaulescent, 10–18 cm tall. Rhizome thesis, much shorter than leaves at fruiting, sparsely pubescent, erect or obliquely ascending, slightly robust, shortly and densely 2-bracteolate at or slightly below middle; bracteoles opposite, noded, with several, scattered, long, yellowish roots. Leaves nu- linear; ca. 1 cm. Sepals oblong or ovate-lanceolate, 5–7 × 3–4 merous, basal; stipules conspicuous, greenish or brownish, sub- mm, apex obtuse or acute, basal auricles subsquare, 1.8–2.5 membranous, adnate to petiole above middle, free part linear- mm, 3-veined, apex shallowly or deeply dentate. Petals obo- lanceolate, ca. 4 mm wide, margin remotely denticulate or en- vate, lateral ones 1.2–1.3 cm × ca. 4.5 mm, slightly bearded, tire; petiole 1–6 cm, wingless, those of lower leaves shorter, anterior one 1.6–1.9 cm (spur included); spur cylindric, 5–7 those of upper leaves longer; leaf blade narrowly triangular or mm, straight, apex rounded. Ovary glabrous; styles slender, ca. oblong, those of lower leaves 2–3 × 1.5–2.5 cm, those of upper 2.5 mm, base slightly geniculate, conspicuously thickened in leaves 4–5 × 1.5–2.5 cm, both surfaces glabrous, veins conspic-

102 VIOLACEAE

uously raised abaxially, base shallowly cordate or truncate, Broad-leaved forests, forest margins, thickets, grasslands on moun- sometimes slightly hastate, margin obtusely crenate, apex ob- tain slopes; 100–600 m. E Jilin, E Liaoning [Japan, Korea, Russia]. tuse. Flowers white, medium-sized, 1.5–1.9 cm; pedicels not or 70. Viola phalacrocarpa Maximowicz, Mélanges Biol. Bull. slightly exceeding leaves, 2-bracteolate at or above middle; Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 9: 726. 1876. bracteoles linear. Sepals ovate-lanceolate or lanceolate, 5–7 mm, apex acuminate, basal auricles short and conspicuous, margin 茜堇菜 qian jin cai narrowly membranous, 3-veined, margin obtusely dentate or Herbs perennial, acaulescent, 6–17 cm tall, dwarf at an- entire, apex truncate. Petals obovate, lateral ones distinctly beard- thesis, conspicuously taller at fruiting. Rhizome erect, short, 3– ed, anterior one broader, apex not emarginate; spur cylindric, 4– 10 mm, stout, covered with white scales, with 2 to several roots; 5 mm, ca. 3 mm in diam., apex rounded. Anthers ca. 2 mm, sub- roots yellow-brown, long, to 18 cm, robust, simple. Leaves bas- equaling appendage of connectives; spur of 2 anterior stamens al, rosulate; outer stipules glaucous, membranous, bladeless, shortly angular, ca. 2.5 mm, apex gradually smaller. Ovary inner ones greenish, over 1/2 adnate to petiole, free part lanceo- glabrous; styles clavate, base slender, slightly geniculate for- late or narrowly lanceolate, apex long acuminate, margin re- ward, gradually thickened upward; stigmas slightly thickened motely shortly fimbriate-denticulate; petiole long, 4–13 cm, on lateral sides and abaxially and narrowly margined, shortly slender, conspicuously winged in upper part, densely puberu- beaked in front, with a smaller stigma hole at tip of beak. Cap- lous when young, later sparsely puberulous; leaf blades of low- sule ellipsoid, 6–9 mm, glabrous, apex often with persistent est leaves often orbicular, those of others ovate or ovate-orbicu- styles. Seeds brownish, ovoid-globose, ca. 1.5 mm, glabrous. lar, 1.5–4.5 × 1.2–2.5 cm, to 6–7 × 5.5–6 cm at fruiting, both Fl. Mar–Apr. 2n = 48. surfaces sparsely or densely (usually on young leaves at anthe- Grassy slopes or wasteland; below 500 m. Jiangsu, Jiangxi, S sis) white puberulous, sometimes purplish abaxially, base slight- Liaoning, Zhejiang [Japan (?naturalized), Korea]. ly cordate but deeply cordate at fruiting, margin crenate, apex obtuse or ± acute. Flowers purple-red, deep purple striate; 69. Viola hirtipes S. Moore, J. Linn. Soc., Bot. 17: 379. 1879. pedicels usually exceeding or subequaling leaves, slender, pu- 毛柄堇菜 mao bing jin cai berulous, rarely glabrous, 2-bracteolate above middle; bracte- oles linear. Sepals lanceolate or ovate-lanceolate, 6–7 mm Herbs perennial, acaulescent and without stolon, 7–15 cm (basal auricles included), margin narrowly membranous, apex tall at anthesis, to 30 cm at fruiting. Rhizome erect, short, less acute, basal auricles 1–2 mm, densely or sparsely puberulous than 1 cm, simple at top, often surrounded by remains of stip- and ciliate, margin usually irregularly shallowly dentate, apex ules; roots usually produced from same place on rhizome, obtuse or truncate. Upper petals obovate, 1.1–1.3 cm × 6–7 oblique or slightly horizontal, white or brownish, 8–15 cm, ro- mm, apex often repand-emarginate; lateral ones oblong-obo- bust, 1.5–3 mm in diam. Leaves 1–4 at anthesis; stipules green- vate, 1.1–1.3 cm × 5–6 mm, distinctly bearded, anterior one ish or glaucous, submembranous, over 1/2 adnate to petiole, 1.7–2.2 mm (spur included), apex retuse; spur tubular, 6–9 free part linear-lanceolate, margin sparsely shortly glandular fim- mm, 1–1.8 mm in diam., straight or curved upward, sometimes briate-denticulate, apex acuminate; petiole equaling or slightly sparsely puberulous, apex rounded. Stamens: appendages of exceeding blade, elongated at fruiting, to 18 cm, densely serice- connectives ca. 1.5 mm; anthers ca. 2 mm; spur of 2 anterior ous; leaf blade oblong-ovate or ovate, 2–7 × 1–4 cm, both sur- stamens long, 6–7 mm, slender, 0.3–0.4 mm in diam. Ovary faces usually glabrous, or white puberulous only in lower part, ovoid-globose, densely puberulous; styles clavate, base genicu- particularly along veins abaxially, base slightly or deeply cor- late, conspicuously thickened upward; stigmas conspicuously date, margin crenate, apex obtuse or ± acute; leaves increasing margined on lateral sides and abaxially, shortly beaked in front, in number at fruiting; leaf blade enlarged to 15 cm × ca. 8 cm. with a large stigma hole at tip of beak. Capsule ellipsoid, 6–8 Flowers purplish, large, 2–3 cm in diam.; pedicels long, slightly mm, with dense short stiff hairs when young, with sparse hairs exceeding or equaling leaves, slender, densely or sparsely white when mature. Seeds red-brown, ovoid-globose, ca. 1.5 mm, ca. puberulous, or sometimes subglabrous in upper part, 2-bracteo- 1 mm in diam. Fl. Apr–May, fr. Jun–Sep. 2n = 24. late at or below middle; bracteoles linear. Sepals oblong-lan- ceolate or narrowly lanceolate, 6–8.5 mm, margin narrowly Grasslands on sunny mountain slopes, thickets, forest margins; membranous, 3- or 5-veined, apex acute, basal auricles 1.5–2 100–600 m. SE Heilongjiang, E Jilin, Liaoning [Japan, Korea, Russia]. mm, ciliate or glabrous, apex obtuse or truncate. Petals obovate, 71. Viola selkirkii Pursh ex Goldie, Edinburgh Philos. J. 6: ca. 1.6 cm × ca. 7.5 mm, lateral ones distinctly bearded, ca. 1.5 324. 1822. × 0.6 cm, anterior one whitish and purple striate at base, 2–2.5 mm (spur included); spur cylindric, 7–9 mm, 1–2 mm in diam., 深山堇菜 shen shan jin cai straight or slightly curved upward, apex rounded. Stamens: Viola selkirkii var. albiflora Nakai; V. selkirkii var. an- appendages of connectives ca. 1.5 mm; anthers 2–2.5 mm; spur gustistipulata W. Becker; V. selkirkii var. brevicalcarata W. of 2 anterior stamens long, 4–5 mm, slender, ca. 0.8 mm in Becker; V. selkirkii var. subbarbata W. Becker; V. selkirkii var. diam. Ovary narrowly ovoid, glabrous or puberulous; styles variegata Nakai. slightly geniculate at base, conspicuously thickened upward; stigmas 2-lobed, conspicuously margined on lateral sides and Herbs perennial, acaulescent and without stolon, 5–16 cm abaxially, shortly beaked in front, with a smaller and upward tall. Rhizome cordlike, occasionally separated, producing vege- stigma hole at tip of beak. Capsule narrowly ellipsoid, glabrous. tative clones. Leaves basal, usually numerous, rosulate; stipules Fl. Apr–Jun, fr. May–Jul. 2n = 24. greenish, ca. 1/2 adnate to petiole, free part lanceolate, margin

VIOLACEAE 103

remotely glandular denticulate, base narrowly and deeply cor- Deciduous mountain forests; below 900 m. E Jilin, Liaoning date, margin obtusely dentate, apex ± acute or obtuse; petiole [Japan, Korea]. 2–7 cm, to 13 cm in fruit, narrowly winged, sparsely white In FRPS (51: 56. 1991), the name Viola monbeigii W. Becker was puberulous or glabrous; leaf blade ovate-cordate or slightly misapplied to this taxon. True V. monbeigii is a synonym of V. be- reniform, 1–5 × 1–3.5 cm at anthesis, ca. 6 × 4 cm at fruiting, lophylla (species no. 66). thinly papery, strigose on the upper surface, glabrous or occa- Viola tokubuchiana var. tokubuchiana occurs in Japan. sionally pubescent below; blades with narrow, deep sinus and lobes touching or slightly overlapping. Flowers long pedicel- 73. Viola violacea Makino, Ill. Fl. Jap. 1(11): t. 67. 1891. late, pedicels 4–7 cm, slightly or not exceeding leaves, gla- 紫背堇菜 zi bei jin cai brous, usually 2-bracteolate at middle; bracteoles linear, 5–7 mm, margin remotely denticulate. Sepals ovate-lanceolate, 6–7 Herbs perennial, acaulescent, 3–8 cm tall, usually gla- mm, margin narrowly membranous, 3-veined, apex acute, basal brous. Rhizome short. Leaves caespitose, ascending, sometimes auricles oblong, ca. 2 mm, apex irregularly emarginate and erect; stipules subulate-lanceolate or subulate-linear, base ad- sparsely ciliate. Petals light blue, purple or white, obovate, all nate to petioles, remotely glandulous-ciliate, membranous; peti- beardless, anterior one 1.5–2 cm (spur included); spur 5–7 mm, ole 2–8 cm; leaf blade abaxially usually purplish, adaxially 2–3 mm in diam., apex rounded, straight or slightly curved up- deep green, sometimes white variegated on veins, narrowly tri- ward. Ovary glabrous; styles clavate, base slightly geniculate angular-ovate, 2–5 × 1–3 cm, both surfaces glabrous or rarely forward, conspicuously thickened in upper part; stigmas flat at pubescent, base cordate, margin shallowly serrate, apex acute. top, narrowly margined on lateral sides, shortly beaked in front, Pedicel 5–8 cm. Flowers red-purple, ca. 1.5 cm in diam. Sepals beak with an upward stigma hole at tip. Capsule ellipsoid, broadly lanceolate, 4–7.5 mm, appendages ca. 1 mm, apex trun- small, 6–8 mm, glabrous, apex obtuse. Seeds numerous, brown- cate or rounded. Petals 8–12 mm; lateral ones glabrous or rarely ish, ovoid-globose, ca. 2 mm, ca. 1.1 mm in diam. Fl. Apr– sparsely bearded; spur usually slightly curved upward, 5–7 mm, May, fr. Jun–Sep. 2n = 12, 24. slender. Ovary ovoid, glabrous. Fl. Mar–Apr, fr. May–Jul.

Mixed needle-leaved and broad-leaved forests, deciduous broad- Forest understories in valleys; below 1000 m. Anhui, Fujian, leaved forests, thickets, stream valleys; 400–1500 m. Hebei, Heilong- Jiangxi, Zhejiang [Japan, S Korea]. jiang, Jilin, Liaoning, Nei Mongol, N Shaanxi [Japan, Korea, Mongolia, 74. Viola macroceras Bunge in Ledebour, Fl. Altaic. 1: 256. Russia; Europe, North America]. 1829. 72. Viola tokubuchiana Makino var. takedana (Makino) F. 大距堇菜 da ju jin cai Maekawa in H. Hara, Enum. Spermatophytarum Japon. 3: 28. 1954. Herbs perennial, acaulescent, 5–14 cm tall, glabrous or sparsely puberulous. Rhizome obliquely ascending or erect, 凤凰堇菜 feng huang jin cai short, with few whitish fibrous roots. Leaves basal; stipules dense in upper part of rhizome, white, ovate-lanceolate, mem- Viola takedana Makino, Bot. Mag. (Tokyo) 21: 57. 1907; branous, ca. 1/2 adnate to petiole, margin entire or sparsely long V. funghuangensis P. Y. Fu & Y. C. Teng. fimbriate, apex acute; petiole 1–8 cm, to 14 cm at fruiting; leaf Herbs perennial, acaulescent, 5–15 cm tall. Rhizome erect blade cordate or ovate-cordate, 1.5–4.5 × 1–3 cm, base shal- or obliquely ascending, short, 0.5–3 cm, slender, with numerous lowly or deeply cordate, usually cuneate and decurrent to peti- white rootlets, sometimes with underground stolons; roots elon- ole, margin shallowly crenate, apex acute, larger at fruiting, to 6 gating, producing new individuals. Leaves several, basal; stip- × ca. 4 cm. Flowers purple-violet or blue-purple, fragrant; pedi- ules nearly 1/2 adnate to petioles, free part lanceolate, 5–10 cels 4–9 cm, 2-bracteolate at middle; bracteoles 4–6 mm. mm, margin remotely denticulate; petiole narrowly winged, 5– Sepals narrowly elliptic or ovate-lanceolate, 5–6 × 2–2.5 mm, apex obtuse, basal auricles short, apex obtuse, glabrous. Petals 11 cm, glabrous, usually purple in lower part, sometimes with 1–1.2 cm, lateral ones bearded; spur 4–6 mm, stout, slightly sparse hairs; leaf blade triangular-ovate to narrowly ovate, 1.8– curved upward or nearly straight. Ovary glabrous; styles clavate, 5 × 1.2–3 cm, larger at fruiting, to 11 × 6 cm, adaxially green to base slightly geniculate, gradually thickened upward; stigmas dark green, sometimes white variegated on veins, both surfaces flat at top, with obtuse margin on lateral sides and abaxially, slightly pubescent or glabrous, base deeply cordate, margin shortly beaked in front, with an upward stigma hole at tip of shallowly crenate-serrate, apex acuminate. Flowers white or beak. Capsule ellipsoid, 7–9 mm, glabrous, apex obtuse. Seeds red-purple; pedicels slender, usually exceeding leaves at anthe- brown or red-brown, large. Fl. Apr–Jun, fr. Aug–Sep. sis, glabrous, 2-bracteolate near middle; bracteoles linear. Se- pals ovate-lanceolate, 6–10 mm; appendages 2–3 mm, apex Alpine meadows, stony slopes; below 1500 m. Xinjiang [Kash- obtuse or shallowly dentate. Lateral petals bearded, anterior one mir, Kazakhstan, Kyrgyzstan, Mongolia, Russia (W Siberia), Tajikistan, Uzbekistan]. 1.4–1.8 cm (spur included), purple striate in lower parts; spur cylindric, 5–6 mm. Ovary glabrous; styles slightly geniculate 75. Viola yunnanfuensis W. Becker, Bull. Misc. Inform. Kew forward at base, gradually thickened upward, stigmas with 1928: 248. 1928. broader margin on two lateral sides and behind, shortly beaked 心叶堇菜 xin ye jin cai in front, with a slightly upward stigma hole at tip. Capsule ellip- soid or ovoid, 7–11 mm, glabrous. Fl. Apr–May, fr. Jun–Aug. Viola bhutanica H. Hara; V. concordifolia C. J. Wang; V.

104 VIOLACEAE

cordifolia W. Becker (1929), not Schweinitz (1822), nor Schur Sepals lanceolate or ovate-lanceolate, 6–8 mm, margin narrow- (1877). ly white membranous, apex acute, basal auricles 1–2 mm, apex irregularly dentate or subentire, ciliate or glabrous. Upper petals Herbs perennial, acaulescent. Rhizome short, robust, 4–5 obovate, 8–11 mm, curved upward, lateral ones oblong-obo- mm in diam., densely noded; roots numerous, whitish, elon- vate, 8–12 mm, glabrous or rarely sparsely bearded, anterior gated, robust. Leaves numerous, basal; stipules short, ca. 1 cm, one 1.4–2.5 cm (spur included); spur 5–9 mm, 1.5–2.5 mm in 1/2–3/4 adnate to petioles, free part spreading; petiole usually diam., apex obtuse and curved upward. Appendage of connec- subequaling blade at anthesis, conspicuously exceeding blade at tives ca. 1.5 mm; anthers 1.5–2 mm; spur of 2 anterior stamens fruiting, very narrowly winged in uppermost part, sparsely pu- ca. 4.5 mm, apex acute. Ovary narrowly ellipsoid, glabrous; berulous or glabrous; leaf blade ovate, broadly ovate, or tri- styles clavate, base conspicuously geniculate, thickened in upper angular-ovate, rarely reniform, 2–8 × 2.5–7 cm, abaxially part; stigmas flat or slightly concave, very rounded or narrowly sparsely puberulous or glabrescent, sometimes purplish, adaxi- margined on lateral sides and abaxially, very shortly beaked in ally spreading white puberulous or glabrescent, base deeply or front, with a smaller stigma hole at tip of beak. Capsule narrow- broadly cordate, margin deeply crenate, apex acute or slightly ly ellipsoid, 5–12 mm, glabrous, apex obtuse, often with persis- obtuse. Flowers white or violet; pedicels not exceeding leaves, tent styles. Seeds numerous, deep brown, ovoid-globose, ca. 2 glabrous, 2-bracteolate near middle; bracteoles linear-lanceo- mm, ca. 1.5 mm in diam., often brown-punctate. Fl. Mar–Apr late. Sepals broadly lanceolate, 5–7 × 1.5–2 mm, apex acumi- and Oct, fr. May–Sep. 2n = 48*. nate, appendages white puberulous or glabrous, ca. 1 mm, apex truncate or slightly dentate. Upper and lateral petals obovate, Grasslands on mountain slopes, streamsides, alongside houses; 1.2–1.4 cm × 5–6 mm, lateral ones conspicuously white below 2800 m. Gansu, Hebei, Heilongjiang, Henan, Hubei, Jilin, Liao- bearded, anterior one narrowly obovate, ca. 1.5 cm (spur inclu- ning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong, Shanxi, Si- chuan [Korea, Russia]. ded), apex emarginate; spur cylindric, 2–3 mm, ca. 2 mm in diam. Ovary conic, glabrous; styles clavate, base slightly geni- 77. Viola japonica Langsdorff ex Candolle, Prodr. 1: 295. culate, thickened in upper part; stigmas flat at top, conspic- 1824. uously margined on lateral sides and abaxially, shortly beaked 犁头草 in front, with a broad stigma hole at tip of beak. Capsule li tou cao ellipsoid, 7–10 mm, glabrous. Fl. Feb–Apr, fr. May–Oct. Viola crassicalcarata Ching J. Wang; V. concordifolia C. Grasslands, thickets; below 3500 m. Guangxi, Guizhou, Sichuan, J. Wang var. hirtipedicellata Ching J. Wang; V. japonica var. S Xizang, Yunnan [Bhutan]. stenopetala Franchet ex H. Boissieu.

76. Viola prionantha Bunge, Mém. Acad. Imp. Sci. St.-Péters- Herbs perennial, acaulescent, rosette-forming. Rhizome bourg Divers Savans 2: 82. 1835. short, 5–7 mm, robust, ca. 5 mm in diam., densely noded; roots numerous, whitish, elongated, robust. Leaves numerous, basal; 早开堇菜 zao kai jin cai stipules narrowly ovate, 2–3 cm × 3–4 mm, ca. 2/3 adnate to petioles, margin remotely ciliate, apex acuminate; petiole usu- Viola prionantha var. sylvatica Kitagawa; V. prionantha ally subequaling blade at anthesis, conspicuously exceeding blade var. trichantha C. J. Wang; V. taishanensis C. J. Wang. at fruiting, very narrowly winged in uppermost part, usually gla- Herbs perennial, acaulescent, 3–10 cm tall at anthesis, to brous; leaf blade ovate, broadly ovate, or triangular-ovate, 3–8 20 cm at fruiting. Rhizome erect, short, 4–20 cm, stout, to 9 × 3–5.5 cm, both surfaces sparsely puberulous or rarely glabres- mm in diam., often surrounded by remains of leaves of previous cent, base deeply or broadly cordate, margin densely crenate, year in upper part; roots several, gray-white, long, robust, usu- apex acute or slightly obtuse. Flowers light purplish or whitish ally produced from lower part of rhizome, straight descending violet; pedicels not exceeding leaves, puberulous or glabrous, 2- or sometimes subhorizontal. Leaves numerous, basal; stipules bracteolate near middle; bracteoles linear-lanceolate, ca. 8 mm. greenish or glaucous, membranous when dry, ca. 2/3 adnate to Sepals broadly lanceolate, 5–7 × ca. 2 mm, apex acuminate, petioles, lower ones 7–9 mm wide, free part linear-lanceolate, basal auricles 2–3 mm, apex truncate or 2-denticulate. Upper 7–13 mm, margin remotely denticulate; petiole 1–5 cm at an- petals oblong-obovate, 1.3–1.5 cm × 5–6 mm, lateral ones ob- thesis, to 13 cm at fruiting, robust, narrowly winged in upper long-obovate, 1.1–1.3 cm × 5–6 mm, sparsely bearded or gla- part, glabrous or puberulous; leaf blade at anthesis oblong- brous, anterior one narrowly obovate, 1.7–2 cm (spur included), ovate, ovate-lanceolate, or narrowly ovate, 1–4.5 cm × 6–20 apex emarginate; spur cylindric, 6–8 mm, 2.5–3.4 mm in diam.; mm, both surfaces glabrous or puberulous, sometimes hairy spur of anterior 2 stamens long, 2–3 mm, slender. Ovary conic, only along veins, base slightly cordate, truncate, or broadly cu- glabrous; styles clavate, base slightly geniculate, thickened in neate, slightly decurrent, margin usually involute when young, upper part; stigmas flat at top, conspicuously margined on lat- densely crenulate, apex obtuse or ± acute; leaf blade conspicu- eral sides and abaxially, shortly beaked in front, with a broad ously enlarged at fruiting, to 10 × 4 cm, triangular-ovate, broad- stigma hole at tip of beak. Capsule ellipsoid, ca. 1 cm. Fl. Mar est near middle, base usually broadly cordate. Flowers purple- and Oct, fr. May–Oct. violet or purplish, light colored and purple striate at throat, Sunny or half shaded places in lowlands; below 1100 m. Anhui, large, 1.2–2 cm in diam., not fragrant; pedicels angled, exceed- Chongqing, Fujian, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, ing leaves, robust, 2-bracteolate near middle; bracteoles linear. Zhejiang [Japan, Korea].

VIOLACEAE 105

78. Viola perpusilla H. Boissieu, Bull. Soc. Bot. France 55: dle; bracteoles nearly opposite, linear, 1–2 mm. Sepals 3–4 × 468. 1908. ca. 1 mm, base acute, margin remotely denticulate-ciliate or glabrous, apex obtuse; basal auricles very short, ca. 0.4 mm, 极细堇菜 ji xi jin cai apex obtuse. Petals violet, oblong-obovate, 5–7 mm, base nar- Herbs perennial, dwarf, acaulescent, 1–5 cm tall. Rhizome rower, apex obtuse, lateral ones glabrous, anterior one slightly erect, brown; roots fibrous, usually undivided. Leaves few, bas- shorter; spur saccate, ca. 2 mm. Anthers ca. 1 mm. Ovary ovoid- al, rosulate; stipules whitish, membranous, 1/3–1/2 adnate to globose, hairy; styles clavate, base conspicuously geniculate, petioles, free part lanceolate, pubescent, apex linear-acuminate; stigma oblique, not margined. Fl. Jun. petiole 0.4–1.5 cm; glabrous; leaf blade ovate or oblong, 0.4– 2.2 cm × 3–8 mm, papery, abaxially glabrous or slightly pu- ● Grassy slopes. Yunnan (Dali). bescent, adaxially pubescent, base rounded or cuneate, slightly decurrent, margin shallowly crenate, apex obtuse. Flowers small; This species is similar to Viola kunawarensis, but has leaf blades pedicels straight, 2–5 cm, longer than leaves, slender, slightly pubescent, margin crenate-dentate, pedicels usually pubescent and much pubescent or nearly glabrous, 2-bracteolate slightly above mid- longer than leaves, and ovary hairy.

2. Viola subg. Dischidium (Gingins) Petermann, Deutschl. Fl. 65. 1846. 二裂花柱亚属 er lie hua zhu ya shu Viola sect. Dischidium Gingins, Mem. Fam. Viol. 1. 1823. Plant with aerial stem. Stipules free, small. Flowers yellow. Anterior petal spurred. Style not beaked at apex; stigmas 2-lobed. About 16 species: Asia, Europe, North America; 11 species (eight endemic) in China. 1a. Rhizome short, stout, erect or obliquely ascending. 2a. Cauline leaves lanceolate and verticillate ...... 79. V. dimorphophylla 2b. Cauline leaves reniform, ovate or triangular, alternate. 3a. Leaf blade pedatipartite ...... 80. V. muliensis 3b. Leaf blade not divided. 4a. Leaf blades reniform or broadly ovate; rhizome very stout, longer, densely noded. 5a. Cauline leaves scattered throughout stem; pedicel 3–4 cm ...... 81. V. szetschwanensis 5b. Cauline leaves usually clustered at top of stem; pedicel 2–3.5 cm ...... 82. V. confertifolia 4b. Leaf blades triangular or ovate; rhizome stouter and shorter. 6a. Leaf blade thickly papery, abaxially glaucous; stipules small, margin entire or remotely dentate ...... 83. V. delavayi 6b. Leaf blade thinly herbaceous, abaxially green; stipules large, margin dentate or shallowly lobed ...... 84. V. urophylla 1b. Rhizome procumbent, slender and long, yellow-brown to whitish, internodes longer, with scattered branched rootlets and nodes. 7a. Leaf blade broadly ovate, triangular-ovate, or ovate-lanceolate. 8a. Leaf blade broadly ovate or triangular ...... 85. V. cameleo 8b. Leaf blade ovate-lanceolate or ovate ...... 86. V. hediniana 7b. Leaf blade reniform or subcordate. 9a. Spur short, 1–2.5 mm ...... 87. V. biflora 9b. Spur longer, 4–6 mm. 10a. Stem filiform; leaf blade basally deeply cordate ...... 88. V. tenuissima 10b. Stem slender; leaf blade basally shallowly cordate ...... 89. V. wallichiana

79. Viola dimorphophylla Y. S. Chen & Q. E. Yang, Bot. J. crenate-serrate, apex obtuse. Cauline leaves (3–)5–7(–10), ver- Linn. Soc. 149: 116. 2005. ticillate, sessile, linear, 8–44 × 1–6.5 mm, both surfaces gla- brous, margin usually entire, sometimes remotely denticulate in 轮叶堇菜 lun ye jin cai upper part, very rarely bifid. Stipules 2, free, leaflike when ma- Herbs perennial, to 30 cm tall, lacking stolons. Rootstock ture, 4–10 mm, nearly indistinguishable from leaves. Flowers to 2.5 cm, ca. 5 mm in diam., vertical, densely articulate, with yellow, axillary, ca. 1.3 cm in diam., yellow; pedicels 1.2–3.5 numerous dark brown fibrous roots. Stems 1–4, 0.2–1 mm in cm, slender, glabrous, 2-bracteolate near apex; bracteoles linear, diam., glabrous, rarely 1-branched. Basal leaves 1 or 2(–4) or very small, ca. 1 mm. Sepals green, linear, 3–4 × 0.25–0.5 mm, withered, long petiolate; petiole 2.5–13 cm, glabrous; leaf blade glabrous, margin entire, apex acuminate; sepal appendages very abaxially purplish, adaxially deep green, broadly cordate, usu- short, glabrous. Petals glabrous, upper ones oblong-spatulate, ally broader than long, 1–4 × 1.5–5 cm, papery, abaxially gla- 7–10 mm, 2.5–3.5 mm wide in upper part, apex rounded; lateral brous, adaxially densely white hispid, base cordate, margin ones oblong-spatulate, 8–11 mm, 3.5–4 mm wide in upper part,

106 VIOLACEAE

apex rounded, at throat with 2 black striae; lower petal broadly V. manaslensis F. Maekawa; V. prattii W. Becker; V. szet- obovate-spatulate, 7–9 mm, 3–4 mm wide in upper part, apex schwanensis var. kangdienensis Chang; V. szetschwanensis var. obtuse, at throat with 6–9 black striae, shortly spurred; spur yel- nudicaulis W. Becker. lowish white, ca. 0.5 × 1.2 mm. Ovary ovoid, glabrous; style Herbs perennial. Stems erect, ca. 25 cm tall, robust, 3- or clavate, base geniculate, 1.6–3 mm; stigma capitate, thick, 4-noded, usually leafless below middle. Stipules narrowly ovate slightly 2-lobed, glabrous. Capsules ovoid or globose, 2–2.5 mm in diam., glabrous. Seeds ovoid, ca. 1.5 × 1 mm. Fl. Jul, fr. to oblong-ovate, ca. 1.3 cm, both surfaces sparsely puberulous Sep. or subglabrous, margin remotely shallowly dentate, apex acu- minate; basal leaves long petiolate; petiole 2–3.5 cm; leaf blade ● Thicket-covered or grassy slopes; 2400–2600 m. NW Yunnan ovate-cordate or broadly ovate, 2–2.5 cm, base cordate, apex (Zhongdian). acuminate; cauline leaf petiole 0.5–1 cm, puberulous, cauline This is a very rare and endangered species. leaf blade broadly ovate, reniform, or suborbicular, 1.5–3 cm wide, abaxially sparsely puberulous, adaxially subglabrous or 80. Viola muliensis Y. S. Chen & Q. E. Yang, Bot. J. Linn. sparsely puberulous along veins, base shallowly cordate, mar- Soc. 149: 365. 2005. gin shallowly crenate, apex mucronate or acuminate. Flowers 木里堇菜 mu li jin cai solitary in leaf axils in upper part, yellow; pedicels erect, much longer than leaves, ca. 3 cm, to 4 cm at fruiting, slender, 2-brac- Herbs perennial, to 28 cm high, lacking stolons. Rootstock teolate at upper part; bracteoles linear. Sepals linear, 4–6 × 0.8– vertical, to 3 cm, densely articulate, with numerous dark brown 1 mm, 3-veined, apex obtuse, basal auricles very short, glabrous fibrous roots. Stems 1–5, 0.5–1 mm in diam., glabrous, very or puberulous, apex truncate. Upper petals oblong, with slender rarely 1- or 2-branched. Leaves broadly ovate or triangular, claw, 1–1.2 cm × ca. 2.5 mm, lateral ones and anterior one very rarely orbicular, 1.5–4.5 × 2–8 cm, papery, both surfaces shorter; spur 2–2.5 mm, apex obtuse. Anthers ca. 1.5 mm, ap- green and glabrous, base truncate to widely cuneate, rarely pendage of connectives 1.5–2 mm; spur of anterior stamens widely cordate, 3-pedatipartite to 3–5 mm from base; middle short, ca. 1 mm. Ovary densely brown punctate; styles genicu- partition rhombic or broadly linear, base cuneate or narrowly late in lower part, thickened in upper part; stigmas 2-lobed, cuneate, apex acuminate, undivided, margin 2- or 3-dentate in lobes auricular, thicker, spreading. Capsule oblong, 0.8–1 cm, upper part, sometimes 2- or 3-fid to 6–7 mm from base, lobes ca. 4 mm in diam., densely brown dotted and sparsely puberu- laxly 2- or 3-incised-lobulate; lateral partitions rarely undivi- lous. Seeds ovoid, ca. 2 × 1 mm. Fl. May–Sep, fr. Jul–Oct. ded, margin 2- or 3-dentate, rarely entire, often unequally 2- or 3-fid, lobes 2- or 3-lobulate. Basal leaves 1–4, sometimes Mountain forests, forest margins, grassy slopes, thickets; 2400– withered, long petiolate; petioles 2.5–14 cm, glabrous. Cauline 4000 m. W Sichuan, Xizang, N Yunnan [Nepal]. leaves similar to basal ones but petioles shorter and blades 82. Viola confertifolia Chang, Bull. Fan Mem. Inst. Biol., n.s., smaller upward. Stipules 2, leaflike, subulate or linear, 4–22 × 1: 238. 1949. 0.5–3 mm, free, glabrous, margin entire, rarely base 1-denticu- late. Flowers axillary, yellow, ca. 1.5 cm in diam.; pedicels 2– 密叶堇菜 mi ye jin cai 4.5 cm, slender, glabrous, near apex 2-bracteolate; bracteoles filiform, very small. Sepals green, narrowly linear, 3–5 × ca. 0.5 Viola biflora Linnaeus var. platyphylla Franchet. mm, glabrous, margin entire, apex acuminate; appendages very Herbs perennial. Rhizome erect, ca. 3.5 cm, ca. 5 mm in short, glabrous. Petals glabrous; upper ones oblong-spatulate, diam., densely noded, with numerous fibrous roots. Stems 1–3, 8–10 mm, 3–4 mm wide in upper part, apex rounded; lateral erect, to 11 cm tall, 2.5–3 mm in diam., glabrous, conspicu- ones oblong-spatulate, 10–12 mm, 2.5–3.5 mm wide in upper ously striate, leafless in middle and lower parts. Basal leaves part, apex rounded, at throat with 2–4 black striae; lower petal few or withered; stipules conspicuous, membranous, margin re- broadly oblong-spatulate, 5–7 mm, 2.5–3 mm wide in upper part, apex obtuse or cuspidate, at throat densely black striate, motely dentate; petiole 5–7 cm, slender, very narrowly winged; shortly spurred; spur yellow, ca. 0.5 mm. Ovary ovoid, gla- leaf blade reniform, abaxially glabrous except along veins, adax- brous; style shortly clavate, base geniculate, 1–1.5 mm; stigma ially sparsely puberulous. Upper leaves clustered at apex of capitate, thick, slightly 2-lobed, glabrous. Capsule ovoid or glo- stem; stipules ovate, 4–5 mm, herbaceous, margin remotely bose, glabrous, ca. 4.5 mm, ca. 2.5 mm in diam. Seeds ovoid, dentate or subentire; petiole ca. 1.5 cm; leaf blade reniform, ca. ca. 0.8 × 1.5 mm. Fl. Jul, fr. Sep. 1.5 × 2.5–4 cm, ± fleshy, both surfaces sparsely puberulous, margin obtusely serrate, apex usually obtuse, rarely mucronate. ● Forest margins, moss-covered rock crevices; ca. 2500 m. SW Flowers few, yellow, long pedicellate; pedicels linear, 2–3.5 Sichuan (Muli). cm, not bracteolate. Sepals linear, ca. 5 mm, 3-veined, margin This is a very rare and critically endangered species. subentire, basal auricles short. Petals obovate-oblong, glabrous, upper ones ca. 7.5 × 2.5 mm, lateral ones and anterior one ca. 8 81. Viola szetschwanensis W. Becker & H. Boissieu, Bull. × 4 mm; spur saccate, ca. 1.5 mm; spur of anterior stamens Herb. Boissier, sér. 2, 8: 742. 1908 [“sze-tschwanensis”]. short. Ovary ovoid-globose, ca. 2 mm, glabrous; styles ca. 1 四川堇菜 si chuan jin cai mm, base conspicuously geniculate, 2-lobed in upper part, lobes narrow and thick. Fl. Jul, fr. Sep. Viola biflora Linnaeus var. acutifolia H. Boissieu; V. biflora var. ciliicalyx H. Boissieu; V. mainlingensis S. Y. Chen; ● Moist places in mountain ravines; 2800–3200 m. NW Yunnan.

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83. Viola delavayi Franchet, Bull. Soc. Bot. France 33: 413. purple striate; spur short, 0.5–0.9 mm, apex obtuse. Stigmas 2- 1886. lobed, lobes spreading, straight, apex obtuse. Capsule ovoid-glo- bose, small. 灰叶堇菜 hui ye jin cai ● Forest margins, meadows, grassy places, streamsides; 1600– Viola boissieui H. Léveillé & Maire; V. delavayi f. depau- 3600 m. Sichuan, W Yunnan. perata Diels; V. delavayi var. villosa W. Becker; V. impatiens H. Léveillé. 1a. Stems, petioles, and leaf blades glabrous ...... 84a. var. urophylla Herbs perennial. Rhizome short, stout, with numerous dark 1b. Stems, petioles, and leaf blades brown fibrous roots. Stems erect, 15–25 cm tall, slender, gla- puberulous ...... 84b. var. densivillosa brous, usually simple, leafless in lower part. Basal leaf usually 1 or absent; petiole to 7 cm; leaf blade abaxially glaucous, adax- 84a. Viola urophylla var. urophylla ially green, ovate, 3–4 × ca. 3 cm, thickly papery, glabrous, 粗齿堇菜(原变种) cu chi jin cai (yuan bian zhong) abaxially sparsely villous at base, base cordate, margin repand- serrate, apex acuminate. Stipules lanceolate, oblong, or ovate, Viola subdelavayi S. H. Huang. 0.5–1.3 mm, lower ones 1–3 × shorter than petiole, upper ones Stems, petioles, and leaf blades glabrous. Fl. May–Jul, fr. subequaling or exceeding petiole, herbaceous, margin entire or Jul–Sep. remotely and large toothed; cauline leaf petiole 0.5–1 cm, gla- brous or rarely villous; cauline leaf blade smaller than those of ● Forest margins on mountains, meadows, shaded and moist basal leaves, broadly ovate or triangular-ovate, base shallowly grasslands along streamsides; 1600–3200 m. SW Sichuan, NW Yunnan. cordate or truncate; upper leaf blade ovate-lanceolate. Flowers 84b. Viola urophylla var. densivillosa C. J. Wang, Acta Bot. in leaf axils at top, yellow, long pedicellate; pedicels exceeding Yunnan. 13: 265. 1991. blades, 1.5–3 cm, 2-bracteolate near top; bracteoles linear. Se- pals linear, ca. 5 mm, glabrous or sparsely puberulous, apex 密毛粗齿堇菜 mi mao cu chi jin cai acute, basal auricles very short, apex truncate. Upper petals Middle and upper parts of stem and petiole densely white obovate, ca. 1.2 cm, lateral ones 0.9–1 cm, anterior one broadly spreading puberulous, stipules and leaf blade sparsely puberu- obovate, 8–9.5 mm, base purple striate; spur very short, only lous on both surfaces, more densely so abaxially on leaf blade, 0.5–1 mm, apex obtuse. Ovary ovoid, smooth, glabrous; styles along veins in particular; leaf blade densely ciliate along mar- slender in lower part, thickened in upper part; stigmas 2-lobed, gin. lobes spreading, straight, broadly ovate, apex rounded. Capsule ovoid or nearly oblong, small, 3–4 mm, subequaling or slightly ● Forest margins, grassy places on mountains, stream valleys; shorter than persistent sepals. Fl. May–Aug, fr. Jul–Oct. 2400–3600 m. Sichuan, SW Yunnan. ● Forest margins on mountains, grassy slopes, moist places in 85. Viola cameleo H. Boissieu, Bull. Herb. Boissier, sér. 2, 1: stream valleys; 1800–2800 m. Guizhou, Sichuan, Yunnan. 1074. 1901. 84. Viola urophylla Franchet, Bull. Soc. Bot. France 33: 413. 阔紫叶堇菜 kuo zi ye jin cai 1886. Viola henryi H. Boissieu var. cameleo (H. Boissieu) Chang. 粗齿堇菜 cu chi jin cai Herbs perennial. Rhizome procumbent, slightly robust, Herbs perennial. Rhizome obliquely ascending, short, covered with scales, with numerous roots. Roots yellowish stout, densely covered with scales, with numerous dark brown when dry, 7–10 cm, with numerous fibrous roots. Stems 15–20 roots. Stems to 22 cm tall, slender, glabrous or pubescent, usu- cm tall, slender, smooth, glabrous. Basal leaves 1 or 2; petiole ally simple, leafless in lower part. Basal leaves usually 1 or 2, long, 6–8 cm, slender; leaf blade broadly ovate or triangular- sometimes absent, long petiolate; leaf blade deep green on both ovate, 1.5–3 × 2–3.5 cm, abaxially puberulous, adaxially usu- sides, deeply cordate, 5–9 × 3–7 cm at base, glabrous or pubes- ally glabrous, base broadly cordate, margin crenate, apex long cent, margin large serrate, apex long acuminate. Cauline leaves: acuminate. Stipules lanceolate, 3–6 mm, herbaceous, margin re- lower stipules nearly ovate, 7–9 mm, much shorter than petiole, motely dentate, apex acute; cauline leaves similar to basal ones, margin remotely and large toothed, upper ones rhombic-ovate but smaller; petiole short, 0.5–2 cm; leaf blade base shallowly or broadly lanceolate, 1.5–2.5 cm, subequaling or longer than cordate or truncate. Flowers solitary in upper leaf axils, golden- petiole, margin remotely and large toothed or shallowly 3- yellow or yellowish, often purple striate; pedicels 2–5 cm, more lobed; petiole 1.5–4 cm; leaf blade slightly glaucous abaxially, than 2 × as long as leaves, slender, 2-bracteolate at upper part; usually triangular-ovate or broadly cordate, 3–5 × 3–6 cm, bracteoles undeveloped, subulate. Sepals very narrow, linear, ca. 0.5 cm × 0.5–0.8 mm, apex acute, basal auricles very short, sometimes sparsely puberulous along veins and margin, base often shallowly dentate. Upper and lateral petals oblong, ca. 1 truncate or broadly cordate. Flowers in axils of cauline leaves, cm × 3–4 mm, not barbate, anterior one obovate; spur tubular, yellow; pedicels 3–4 cm, slightly shorter than leaves, 2-bract- 3–4 mm, ca. 1 mm in diam., slightly slender, straight or slightly eolate below flower; bracteoles undeveloped, linear. Sepals lin- curved. Ovary ovoid, smooth, glabrous; styles 2-lobed in upper ear, ca. 4 mm, apex acute, basal auricles short, 3-veined, apex part, lobes oblong, apex obtuse. Fl. May–Jul, fr. Aug–Sep. truncate. Upper petals obovate, ca. 1.1 cm, lateral ones spat- ulate, ca. 1 cm, anterior one broadly obovate, ca. 8 mm, base ● Thickets; 1800–3800 m. W Hubei, Sichuan, NW Yunnan.

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86. Viola hediniana W. Becker, Beih. Bot. Centralbl., Abt. 2, forest margins, rocks and rock crevices; 2500–4300 m. Gansu, Hebei, 34: 262. 1916. Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan, Xinjiang, Xizang, Yun- 紫叶堇菜 zi ye jin cai nan [Bhutan, NE India, Indonesia (N Sumatra), Japan, Kashmir, Korea, Malaysia, Mongolia, Myanmar, Nepal, Russia; Europe, North America]. Herbs perennial. Rhizome procumbent or obliquely as- cending, whitish, short, slightly robust, with numerous rootlets. 1a. Leaf blade reniform or broadly ovate, Stems erect, 25–30 cm tall, slender, glabrous, usually simple, usually larger ...... 87a. var. biflora leafless in lower part. Basal leaves 1 or 2 or absent, long petio- 1b. Leaf blade orbicular or nearly reniform, late; lower cauline leaves shortly petiolate, upper one subsessile smaller, 1–1.5 × 1–1.5 cm ...... 87b. var. rockiana or sessile; stipules free, ovate-lanceolate or lanceolate, small, 3– 87a. Viola biflora var. biflora 5 mm, herbaceous, margin remotely dentate; leaf blade dark green adaxially, ovate-lanceolate or ovate, 3–7 × 1.5–2.5 cm, 双花堇菜(原变种) shuang hua jin cai (yuan bian zhong) abaxially glabrous, adaxially sparsely puberulous or subgla- Viola biflora var. acuminata Maximowicz; V. biflora var. brous, base shallowly cordate or rounded, margin crenate, apex hirsuta W. Becker; V. biflora var. nudicaulis W. Becker; V. long acuminate. Flowers solitary in upper leaf axils, yellow; biflora var. valdepilosa Handel-Mazzetti; V. chingiana W. pedicels erect, slightly shorter than leaves, slender, 2-bracteo- Becker; V. kanoi Sasaki; V. nudicaulis (W. Becker) S. Y. Chen; late near middle; bracteoles subulate. Sepals linear, ca. 5 × 0.5– V. schulzeana W. Becker; V. tayemonii Hayata; V. wallichiana 1 mm, apex acute, basal auricles very inconspicuous, apex trun- subsp. brevicornis W. Becker. cate. Upper and lateral petals oblong, 8–12 × 3–4 mm, lateral ones inside not barbate, anterior one triangular-obovate, ca. 1.3 Stems usually 10–25 cm tall. Leaf blade reniform or cm (spur included), base dark yellow; spur cylindric, 5–6 mm, broadly ovate, 1–3 × 1–4.5 cm, base deeply cordate or cordate, 1.5–2 mm in diam.; spur of anterior stamens cylindric, 2–2.5 margin obtusely dentate, apex obtuse; spur 1–2.5 mm. Fl. May– mm, slender. Ovary smooth, glabrous; styles slightly geniculate Jul, fr. Jul–Oct. 2n = 12*. at base, conspicuously thickened upward, 2-lobed in upper half, Alpine or subalpine meadows, thickets, forest margins, rock lobes ovate, apex acute. Capsule ellipsoid, ca. 6 mm, glabrous. crevices; 2500–4000 m. Gansu, Hebei, Heilongjiang, Henan, Jilin, Fl. Apr–Jun, fr. Jul–Sep. Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Taiwan, Xinjiang, Xizang, Yunnan [Bhutan, NE India, Indonesia (N ● Mountain forests, forest margins, grassy slopes, moist rock Sumatra), Japan, Kashmir, Korea, Malaysia, Mongolia, Myanmar, crevices; 1500–3500 m. W Hubei, Sichuan. Nepal, Russia; Europe, North America].

87. Viola biflora Linnaeus, Sp. Pl. 2: 936. 1753. 87b. Viola biflora var. rockiana (W. Becker) Y. S. Chen, 双花堇菜 shuang hua jin cai comb. et stat. nov. 圆叶小堇菜 Herbs perennial. Rhizome erect or obliquely ascending, yuan ye xiao jin cai slender or slightly robust, noded, with numerous rootlets. Stems Basionym: Viola rockiana W. Becker, Repert. Spec. Nov. 2 or several fasciculate, erect or obliquely ascending, 10–25 cm Regni Veg. 21: 236. 1925; V. jizushanensis S. H. Huang. tall, slender, 3–5-noded, usually glabrous or sparsely puberu- lous when young. Basal leaves 2 to several; petiole 4–8 cm; leaf Stems usually 2(or 3), slender, to 10 cm tall, 2-noded, gla- blade reniform, broadly ovate, or suborbicular to orbicular, 1–3 brous, with leaves only in lower part. Leaf blade orbicular or × 1–4.5 cm, abaxially glabrous, adaxially sparsely puberulous, ovate-orbicular, ca. 1 × 1 cm, base shallowly cordate or sub- sometimes puberulous on both surfaces, base cordate or sub- truncate, margin shallowly repand-crenate. Spur shallowly sac- truncate, margin obtusely dentate or shallowly repand-crenate, cate, 1–1.5 mm. Fl. Jun–Jul, fr. Jul–Aug. apex obtuse. Stipules free, ovate or ovate-lanceolate, 3–6 mm, ● Thickets, alpine or subalpine grassy slopes, rocks; 2500–4300 margin entire or remotely denticulate, apex acute; cauline leaves m. Gansu, Qinghai, Sichuan, E Xizang, Yunnan. shortly petiolate; petiole glabrous or puberulous; leaf blade smaller than basal ones. Flowers yellow, sometimes whitish at 88. Viola tenuissima Chang, Bull. Fan Mem. Inst. Biol., n.s., 1: 234. 1949. late anthesis; pedicels 1–6 cm, slender, 2-bracteolate at upper part; bracteoles lanceolate. Sepals linear-lanceolate or lanceo- 纤茎堇菜 xian jing jin cai late, 3–4 mm, apex acute, basal auricles very short, margin mem- branous, glabrous or shortly ciliate in middle and lower parts. Herbs perennial, small, 5–7 cm tall. Rhizome short, dense- Petals oblong-obovate, 6–8 mm, purple veined, lateral ones in- ly covered with few scales. Stems greenish when dry, filiform, side not barbate, anterior one ca. 1 cm (spur included); spur slender, glabrous. Basal leaf usually 1, long petiolate; stipules narrowly ovate, 2–2.5 mm, margin subentire or sometimes re- shortly cylindric, 0.5–2.5 mm; spur of anterior stamens shortly motely and inconspicuously serrate; cauline leaf petiole linear, triangular. Ovary glabrous; styles clavate, base slightly genicu- long, to 5 cm, slender; cauline leaf blade abaxially glaucous, late, deeply 2-lobed in upper half, lobes obliquely spreading, adaxially green, cordate, 1–1.5 × 1–1.5 cm, both surfaces with a conspicuous stigma hole in between lobes. Capsule ob- sparsely white puberulous or subglabrous, base deeply cordate, long-ovoid, 4–7 mm, glabrous. lateral auricular lobes rounded and often overlapping, margin Alpine or subalpine meadows and grassy slopes, thickets, forests, remotely and large serrate, apex shortly or inconspicuously

VIOLACEAE 109

acuminate. Flowers solitary in upper leaf axils, yellow; pedicels very sparsely puberulous. Basal leaves 2–4; petiole long, 2.5–6 filiform, 2–3.5 cm, slender, 2-bracteolate above middle; brac- cm, slender; leaf blade reniform or suborbicular, 8–15 mm × teoles linear, small. Sepals very narrow, linear, ca. 5 × 0.6–1 1.5–2.5 cm, both surfaces glabrous, puberulous only along mar- mm, basal auricles very short, glabrous, margin narrowly mem- gin, base cordate, margin crenate, apex rounded. Cauline leaves branous, apex truncate. Upper petals oblong, ca. 1 × 0.3 cm, smaller; stipules free, narrowly ovate to lanceolate, ca. 3 mm, lateral ones not barbate, slightly longer than upper ones, anteri- submembranous, apex acuminate; petiole shorter, 2–10 mm; leaf or one narrowly obovate, ca. 1 cm × 4–7 mm, base purple stri- blade reniform or nearly cordate, base shallowly cordate or trun- ate; spur 3–5 mm, 1–1.5 mm in diam., apex small and obtuse. cate, apex acute. Flowers in upper leaf axils, yellow; pedicels Ovary glabrous, red punctate; styles 2–2.5 mm, base geniculate, 2–3 cm, slender, bracteolate at upper part; bracteoles reduced, apex 2-lobed, lobes obtuse. Capsule subglobose, ca. 4 mm in membranous. Sepals narrow, subulate, 4–6 × 0.8–1 mm, apex diam., red punctate. Fl. Jun–Jul, fr. Aug–Sep. acute, basal auricles inconspicuous, margin narrowly membra- nous, glabrous. Upper and lateral petals oblong-obovate, 7–9 × ● Mountain forests, rock crevices in shaded and moist places; 2.5–3 mm, anterior one broadly obovate, ca. 1.4 cm (spur in- 2300–3300 m. N Guizhou, W Sichuan. cluded), ca. 7 mm wide, apex shallowly 2-dentate; spur tubular, 89. Viola wallichiana Gingins in Candolle, Prodr. 1: 300. 4–6 mm, ca. 0.8 mm in diam., apex acute; spur of anterior sta- 1824. mens angular, ca. 3 mm. Ovary ovoid-globose, glabrous; styles clavate, base conspicuously geniculate, 2-lobed in upper part, 西藏细距堇菜 xi zang xi ju jin cai lobes spreading, thick. Capsule ovoid-globose, ca. 3 mm, gla- Herbs perennial. Rhizome short, robust, densely noded, brous. Fl. and fr. Jul–Aug. covered with scales, with numerous rootlets. Stems 1–4, 5–10 Rock crevices in forest understories; ca. 2900 m. Xizang [India cm tall, slender, 4–6-noded, leafless in lower part, glabrous or (Darjeeling, Sikkim), Nepal].

3. Viola subg. Chamaemelanium (Gingins) Juzepczuk in Schischkin & Bobrov, Fl. URSS 15: 446. 1949. 须毛柱头堇菜亚属 xu mao zhu tou jin cai ya shu Viola sect. Chamaemelanium Gingins, Mem. Fam. Viol. 1. 1823. Plant with aerial stem. Stipules free, small. Flowers yellow, rarely white or with violet or brown upper petals. Anterior petal spurred. Style gradually globose, incrassate and not beaked at apex; stigmas on front side of style incrassation, on hemispheric projection. About 26 species: NE Asia, North America; three species in China. 1a. Rhizome procumbent, slender, long, yellow-brown to whitish, internodes longer, with scattered branched rootlets and nodes; flowers larger, 1.5–2 cm ...... 92. V. muehldorfii 1b. Rhizome short, stout, erect or obliquely ascending; flowers smaller, 1–1.5 cm. 2a. Lowest cauline leaf above middle of stem; pedicel shorter, 1–3 cm ...... 90. V. orientalis 2b. Lowest cauline leaf below middle of stem; pedicel longer ...... 91. V. acutifolia 90. Viola orientalis (Maximowicz) W. Becker, Beih. Bot. eoles usually opposite, sometimes alternate, ovate, small, 1–2 Centralbl., Abt. 2, 34: 265. 1916. mm. Sepals lanceolate or oblong-lanceolate, 5–7 mm, apex acute, basal auricles suborbicular, short. Petals obovate, upper 东方堇菜 dong fang jin cai and lateral petals curved toward outside, upper ones dark purple Viola uniflora Linnaeus var. orientalis Maximowicz, striate, lateral ones inside conspicuously barbate, anterior one Enum. Pl. Mongolia, 81. 1889; V. conferta (W. Becker) Nakai; shorter, 1–1.5 cm (spur included); spur saccate, short, 1–2 mm; V. orientalis var. conferta W. Becker; V. xanthopetala Nakai. spur of anterior stamens ca. 0.5 mm wide. Ovary glabrous; Herbs perennial. Rhizome usually erect, sometimes styles slightly straight at base, gradually thickened upward; stig- obliquely ascending, 1–3 cm, robust, ca. 5 mm in diam., dense- mas capitate, with several series of long beards on lateral sides. ly noded; roots numerous, white to yellowish brown, to 10 cm, Capsule greenish, ellipsoid or oblong, 7–12 mm, often dark pur- with numerous filiform rootlets in lower part. Stems erect, 6–10 ple punctate. Seeds white to brownish, ovoid-globose, ca. 2 cm tall, usually puberulous. Stipules small, adnate to petiole mm. Fl. Apr–May, fr. Jun–Jul. 2n = 12. only at base, free part ovate, 1–2 mm, margin entire or remotely Sparse forests on mountains, forest margins, thickets, grasslands serrulate; basal leaf petiole 3–10 cm; blade ovate, broadly so, or on mountain slopes; 100–1100 m. Heilongjiang, E Jilin, Liaoning, E elliptic, 2–4 × 1.5–3 cm, abaxially puberulous, adaxially sub- Shandong [Japan, Korea, Russia]. glabrous, base cordate, sometimes subtruncate, margin obtusely 91. Viola acutifolia (Karelin & Kirilov) W. Becker, Beih. Bot. serrate, apex acute; cauline leaves 3(or 4), upper 2 very shortly Centralbl., Abt. 2, 34: 263. 1916. petiolate or subsessile, subopposite, lower ones shortly petio- 尖叶堇菜 late, above middle of stem. Flowers yellow, ca. 2 mm in diam., jian ye jin cai usually 1–3 in cauline leaf axils; pedicels 1–3 cm, white puber- Viola biflora Linnaeus var. acutifolia Karelin & Kirilov, ulous, sometimes glabrous, 2-bracteolate at upper part; bract- Bull. Soc. Imp. Naturalistes Moscou 15: 163. 1842.

110 VIOLACEAE

Herbs perennial. Rhizome obliquely ascending or erect, and white rootlets. Stems erect, simple, 20–60 cm tall, spread- with numerous yellow-brown rootlets. Stems 1–3, 10–25 cm ing white villous or subglabrous in lower part, usually with 1 tall, usually flexible, glabrous or spreading puberulous. Basal small, white, ovate, membranous scale at base. Basal leaves 1– leaves 1 or 2, rarely 5, usually withered after anthesis; leaf 3, petiole to 10 cm; leaf blade cordate or reniform, both sur- blade cordate or reniform, 3–5 × 3.5–5.5(–6) cm, both surfaces faces sparsely puberulous, more densely puberulous on veins subglabrous or sparsely puberulous, base deeply cordate, apex abaxially, base cordate, margin serrate and ciliate, apex mucro- obtuse or mucronate. Stipules adnate to petiole only at base, nate. Cauline leaves usually 3, rarely 4; stipules free, opposite, free part ovate-lanceolate, 4–6 mm, margin entire or remotely ovate, margin entire or glandular serrate, apex acute; petiole of fimbriate-dentate, apex acute; cauline leaf only 1 in lower part lower one ca. 3 cm, densely minutely villous, blade of lower of stem, long petiolate, without flowers in leaf axils; leaves at one orbicular-cordate, ca. 4 × 4.5 cm, base broadly cordate, apex apex of stem subverticillate, shortly petiolate; leaf blade broadly acuminate; upper two short petiolate or subsessile, leaf blade of ovate, 5–7 × 3–6 cm, both surfaces sparsely puberulous, base upper two at apex of stem, subopposite, ovate, both surfaces cordate, margin remotely serrate, apex acuminate. Flowers 1–3, white puberulous, base shallowly cordate, apex acuminate or usually 2 in leaf axils at apex of stem, yellowish, 1–2 cm in acute. Flowers in second leaf axil at apex of stem, golden-yel- diam.; pedicels shorter than leaves, 2-bracteolate above middle; low, 1.5–2 cm; pedicels glabrous, 2-bracteolate in upper part; bracteoles linear. Sepals linear-lanceolate or linear, 6–8 mm, bracteoles broadly ovate. Sepals narrowly ovate or lanceolate, apex ± acute, basal auricles very short, 3-veined. Petals obo- ca. 8 mm, usually glabrous, 3-veined, margin entire, basal auri- vate, purple striate, anterior one shorter, 1.3–1.5 cm (spur in- cles short, ca. 0.3 mm. Petals obovate, purple striate, lateral cluded); spur saccate, very short, slightly curved upward. Cap- petals inside barbate at base, anterior one subspatulate, 1.5–2 sule oblong, 6–12 mm, glabrous. Fl. May–Jun, fr. Jun–Jul. cm (spur included); spur ca. 2 mm, stout, apex obtuse; spur of Alpine or subalpine meadows, grasslands on mountain slopes, anterior stamens undeveloped, short, broad. Ovary glabrous; forest margins; 1000–2400 m. Xinjiang [Russia]. styles gradually thickened upward; stigmas capitate, with a series 92. Viola muehldorfii Kiss, Bot. Közlem. 19: 92. 1921 of white beards on each lateral side, shortly beaked in central [“mühldorfi”]. part, with a rounded stigma hole at tip of beak. Capsule ellip- soid, ca. 7 mm, apex ± acute, usually with persistent style. Seeds 大黄花堇菜 da huang hua jin cai glaucous, globose, ca. 2 mm in diam., with white appendage. Viola lasiostipes Nakai. Fl. May–Aug, fr. Jul–Sep. Herbs perennial. Rhizome procumbent, yellow-brown to Mixed needle-leaved and broad-leaved forests, forest margins, whitish, long, slender, with small, dry, white membranous scales, streamsides; 300–500 m. Heilongjiang [Korea, Russia].

4. Viola subg. Melanium (Gingins) Petermann, Deutschl. Fl. 65. 1846. 美丽堇菜亚属 mei li jin cai ya shu Viola sect. Melanium Gingins, Mem. Fam. Viol. 1. 1823. Plant with aerial stem (in China plants). Stipules free, large, often leaflike, deeply lobed. Style globose and not beaked at apex; stigmas on front side of style incrassation. Anterior petal broadly cuneate, spurred. Between 80 and 100 species: N Africa, Asia, Europe, occasionally in North and South America; three species (two introduced) in China. 1a. Stem shorter, usually 1-flowered; leaf blade orbicular-ovate or oblong-ovate, leaves usually clustered ...... 93. V. altaica 1b. Stem longer, usually many flowered; blade of upper leaves oblong-lanceolate, leaves usually scattered, alternate. 2a. Corolla flat; petals variously marked with yellow, white, or purple, ca. 3 × as long as sepals; blade of lower and median leaves basally rounded or cordate ...... 94. V. tricolor 2b. Corolla funnel-shaped; petals ivory or pale yellow, shorter to slightly longer than sepals; blade of lower and median leaves basally cuneate ...... 95. V. arvensis 93. Viola altaica Ker Gawler, Bot. Reg. 1: t. 54. 1815. glabrous. Sepals oblong-lanceolate, 8–13 × 3–4 mm, margin us- ually remotely denticulate, apex ± acute, basal auricles broader, 阿尔泰堇菜 a er tai jin cai often denticulate. Upper petals subovate-orbicular, lateral ones Herbs perennial, 4–15 cm tall. Rhizome slender, branched. and anterior one conspicuously purple-black striate, lateral ones Stems very short, densely covered with numerous leaves usu- inside usually slightly barbate at base; spur of anterior petal 3–4 ally clustered. Stipules ovate or oblong, 1–2.6 cm × 4–11 mm, mm, slightly exceeding basal auricles of sepals, usually slightly pinnatifid, terminal lobe larger, oblong-ovate, lateral lobes 2 or curved upward. Capsule oblong-ovoid. Fl. Jun–Aug, fr. Sep. 2n 3 on each side, lanceolate or oblong-lanceolate, margin sparsely = 54. puberulous; petiole usually longer than leaf blade; leaf blade orbicular-ovate or oblong-ovate, 1.2–2 × 1–1.4 cm, both sur- Alpine and subalpine meadows, forests on mountain slopes, grass- faces subglabrous or sparsely puberulous, base ± cuneate, mar- lands, among mosses and lichens; 1500–4000 m. Xinjiang [Kazakh- gin rounded-dentate, apex obtuse. Flowers usually solitary, yel- stan, Kyrgyzstan, Mongolia, Russia (Siberia); SW Asia (Caucasus), low or blue purple, large, 2–4.5 cm in diam.; pedicels 5–16 cm, SE Europe (Ukraine)].

VIOLACEAE 111

94. Viola tricolor Linnaeus, Sp. Pl. 2: 935. 1753. Widely cultivated in China [native to Europe]. 三色堇 san se jin 95. Viola arvensis Murray, Prodr. Stirp. Gött. 73. 1770. 野生堇菜 Viola tricolor var. hortensis Candolle. ye sheng jin cai Herbs annual or biennial, 5–50 cm tall. Stems erect to as- Herbs annual, biennial, or perennial, 10–40 cm tall. Stems cending, simple or with ascending branches from base, very erect or slightly oblique, slightly robust, angled, simple or nu- sparsely puberulous. Stipules 2–40 mm, pinnatipartite, terminal merously branched. Basal leaf blade narrowly ovate or lanceo- segments largest, often leaflike, lateral ones linear to narrowly late, long petiolate; stipules large, leaflike, deeply pinnatifid, 1– lanceolate; leaf blade 0.5–6.5 × 0.2–2.2 cm, sparsely puberu- 4 cm; petioles of upper leaves longer, those of lower leaves lous, especially so along margin and along veins abaxially, mar- shorter; cauline leaf blade ovate, oblong-orbicular, or oblong- gin crenate or serrate; lower leaves ovate or orbicular-ovate, lanceolate, base rounded, margin remotely crenate, apex round- base rounded, petioles equaling or exceeding blades; middle ed or obtuse. Flowers solitary in leaf axils, usually 3-colored leaves oblong-ovate or oblong-elliptic, base cuneate, decurrent (purple, white, and yellow), large, 3.5–6 cm in diam., with 3–10 into a short petiole; upper leaves oblong-lanceolate, with indis- flowers per stem; pedicels slightly robust, 2-bracteolate at upper tinct petioles. Flowers few to numerous; pedicels 2.5–11 cm, part; bracteoles opposite, ovate-triangular, very small. Sepals glabrous, 2-bracteolate; bracteoles lanceolate. Sepals oblong- green, oblong-lanceolate, 1.2–2.2 cm × 3–5 mm, margin nar- lanceolate, 4–15 mm, apex acute; basal auricles 1–4 mm, ir- rowly membranous, apex acute, basal auricles developed, 3–6 regularly toothed. Corolla funnelform; petals yellow or ivory, mm, margin irregular. Corolla flat; upper petals deep purple- obviously shorter than or nearly equaling sepals, 6–14 mm, up- violet, lateral petals and anterior one 3-colored, purple-striate, per ones pale, anterior one including spur 5.5–14 mm; spur 1–4 lateral ones inside densely barbate at base; spur of anterior petal mm, not exceeding appendages, straight. Capsule 6–10 mm. smaller, 5–8 mm. Ovary glabrous; styles short, base conspic- Seeds oblong-ovoid, 1.5–1.8 mm. Fl. Apr–Sep, fr. Jun–Sep. 2n uously geniculate; stigmas enlarged, globose, with a large stig- = 34. ma hole in front. Capsule ellipsoid, 8–12 mm, glabrous. Fl. Weeds in exposed places, abandoned fields, or cultivated. Taiwan Apr–Jul, fr. May–Aug. 2n = 26. [native to N Africa, SW Asia, and Europe].

5. Viola subg. Erpetion (Sweet) Y. S. Chen, stat. nov. 无距堇菜亚属 wu ju jin cai ya shu Basionym: Erpetion Sweet, Brit. Fl. Gard. 2: t. 170. 1826; Viola sect. Erpetion (Sweet) W. Becker. Stipules free, brownish green, small. Flowers white to violet. Anterior petal shallowly saccate but not spurred. One species: Australia; introduced in China. 96. Viola hederacea Labillardière, Pl. Nov. Holl. 1: 66. 1805. diam., sometimes to 15 mm, scentless or faintly scented. Se- pals green, ovate to linear-lanceolate, 1.5–5 mm, glabrous or 常春藤叶堇菜 chang chun teng ye jin cai sparsely puberulous, basal appendages short. Petals white or pale to dark violet, 2–10 mm, anterior one shallowly saccate, Herbs perennial, stoloniferous. Stem erect, short. Leaves spurless, lateral ones somewhat falcate, bearded inside or gla- clustered; stipules linear-lanceolate, 1.5–8 mm, margin glan- brous. Stamens to 2 mm; filaments 0–0.3 mm. Capsule ovoid, dular-denticulate; petiole 0.5–6 cm, sometimes to 12 cm; leaf 4–6 mm. Seeds white, brown, or black, ovoid, 1.5–2 mm. Fl. blade ovate, reniform, or suborbicular, glabrous or puberulous, Feb–Apr, fr. May–Sep. base cuneate, truncate, or slightly cordate, margin entire to large repand or crenate. Flowers with scape to 10 cm high, glabrous Weed communities under trees and shrubs, grasslands, introduced. or puberulous, bracteolate near middle. Flowers 3–10 mm in Hong Kong [native to Australia].